EVALUATION OF FLORIDA BAY SEAGRASS AFTER HURRICANE …
Transcript of EVALUATION OF FLORIDA BAY SEAGRASS AFTER HURRICANE …
EVALUATION OF FLORIDA BAY SEAGRASS
AFTER HURRICANE IRMA
by
AUBREY LYNN LORIA
MICHAEL STEINBERG, COMMITTEE CHAIR
JULIA CHERRY
NICHOLAS MAGLIOCCA
A THESIS
Submitted in partial fulfillment of the requirements
For the degree of Master of Science
in the Department of Geography
in the Graduate School of
The University of Alabama
TUSCALOOSA, ALABAMA
2019
Copyright Aubrey Lynn Loria 2019
ALL RIGHTS RESERVED
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ABSTRACT
When Hurricane Irma made US landfall on the Florida Keys on September 10, 2017, the
category 4 storm became the first major hurricane to hit the state since 2005. This study
investigated the impacts that Irma had on seagrasses in Florida Bay – an estuary at the southern
outlet of the Florida Everglades that is crucial to the environment and economy of South Florida
but that is threatened by anthropogenic impacts on water quality. A massive seagrass die-off in
2015 left the bay with hypersaline water and sulfide-saturated sediment that was held in place by
the bay’s network of carbonate mud banks. The promise of rainfall and storm surge from a major
hurricane offered a chance to refresh the bay and begin the process of recovering from the die-
off. By comparing seagrass density and community composition from after the 2015 die-off to
after Irma, I found that seagrass cover has become more uniform and that species richness has
increased throughout the study area. Seagrass density was lost in some shallow sections of the
study area, but pioneer species Halodule wrightii had begun colonizing the disturbed areas,
indicating that water quality had changed enough to support the less salt-tolerant species. The
positive changes in seagrass density, distribution, and diversity after Hurricane Irma indicate that
the storm may have flushed out the hypersaline water left after the 2015 die-off that had been
limiting seagrass recovery for two years.
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DEDICATION
Dedicated to the memory of my hero, teacher, and grandfather, Don, who meant to make
me an artist but accidentally made me a scientist. Don passed away two weeks after I completed
the field research for this thesis, but the love and pride he showed for his grandchildren has
continued to encourage me in the months since.
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LIST OF ABBREVIATIONS AND SYMBOLS
D Density using the Braun-Blanquet Cover-Abundance scale
PSU Practical Salinity Units
RKB Rabbit Key Basin
TKB Twin Key Basin
< Less than
= Equal to
≤ Less than or equal to
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ACKNOWLEDGMENTS
This thesis simply could not have been done without all the help I received from my
incredible mentors, family, and support system. First, I thank Dr. Michael Steinberg (Committee
Chairman), for accepting my request to join his lab and encouraging me to revisit the work of his
former graduate student, Cynthia “Jo” McGinnis. Thank you to Jo for providing the groundwork
that makes this comparison so interesting and for making a trip to come speak with me at the
University of Alabama while I was putting together my thesis proposal.
Thank you to Everglades National Park (ENP), South Florida Natural Resources Center
(SFNRC), and the Florida Bay Interagency Science Center (FBISC) for providing logistical
support for this thesis. More specifically, thank you to Matt Patterson of the SFNRC for not only
coordinating this thesis with NPS and FBISC, but also for making sure I never had any question
go unanswered, I never went without any equipment that he could provide, and I never felt alone
when I was so far from home. Thank you to Dr. Jed Redwine for advice on several
methodologies. Thank you to Troy Mullins for helping me acquire GIS datasets.
Thank you to Eduarda and Mima for providing me with free room and board (respectively) for
nine weeks while I conducted my research in South Florida. It was wonderful getting to spend
that time with both of you.
Finally, thank you to my parents for my life, my sister for my ambition, and Stirton and
Lita for my courage. Thank you all for getting me here.
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CONTENTS ABSTRACT ................................................................................................................................ ii
DEDICATION ........................................................................................................................... iii
LIST OF ABBREVIATIONS AND SYMBOLS ...................................................................... iv
ACKNOWLEDGMENTS .......................................................................................................... v
LIST OF TABLES ................................................................................................................... viii
LIST OF FIGURES ................................................................................................................... ix
CHAPTER 1 INTRODUCTION ................................................................................................ 1
CHAPTER 2 BACKGROUND .................................................................................................. 5
2.1 Florida Bay Ecosystem ..................................................................................................... 5
2.2 Florida Bay Hydrology ..................................................................................................... 7
2.3 Plants Studied.................................................................................................................. 11
2.4 Hurricane Effects ............................................................................................................ 13
CHAPTER 3 MATERIALS AND METHODS ....................................................................... 17
3.1 Study Sites ...................................................................................................................... 17
3.2 Sampling Method ............................................................................................................ 19
3.3 Spatial Analysis .............................................................................................................. 22
CHAPTER 4 RESULTS ........................................................................................................... 24
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CHAPTER 5 DISCUSSION ..................................................................................................... 38
CHAPTER 6 CONCLUSION................................................................................................... 41
REFERENCES ......................................................................................................................... 42
APPENDIX ............................................................................................................................... 47
Accuracy Assessment ........................................................................................................... 47
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LIST OF TABLES
Table 1(a-b): Braun-Blanquet assessment of seagrass density (D) for all samples, separated by
species and basin. 1a contains pre-Irma data and 1b contains post-Irma. Adapted from
Fourqurean et al. (2001). ...............................................................................................................28
Table 2: Root Mean Square Error for each map by basin (when applicable) ................................47
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LIST OF FIGURES
1. Everglades National Park, Florida, USA. Park extent (green outline) and Florida Bay
extent (black dashed line) are shown .........................................................................................4
2. Annual mean volume transports (m3 s−1) for measured channels in Florida Bay (red
arrows and numbers), through the tidal passes between the Florida Keys (blue arrows
and numbers), and from mean river discharge (black arrows and numbers)
(Lee et al. 2016) .......................................................................................................................10
3. Thalassia testudinum (top), Halodule wrightii (middle), and Syringodium filiforme
(bottom). Illustrations are by Phillips and Menez (1988) and photographs are by Ken
Dunton (University of Texas). .................................................................................................11
4. Hurricane Irma's track in relation to Florida Bay and South Florida .......................................13
5. A wrack line composed entirely of dried seagrass, mostly Thalassia testudinum, left by
Hurricane Irma. This photo was taken over 300 meters from Flamingo Beach, FL. ..............15
6. Florida Bay salinity (PSU) one month before Hurricane Irma, one month after, and five
months after (SFWMD 2018). .................................................................................................16
7. Twin Key and Rabbit Key Basins and their position within Florida Bay. ..............................18
8. Bathymetry of Rabbit Key and Twin Key Basins. Contour interval: 0.25 m. Digital
Elevation Model provided by National Centers for Environmental Information (NCEI).
Vertical error estimated by NCEI to be no more than 15 cm ..................................................19
9. Location of sampling sites in RKB and TKB. .........................................................................20
10. Total seagrass cover defined by broad cover categories. Bare sediment is 0% cover;
Sparse cover is <10%; Moderate cover is 10-50%; dense cover is >50%. ..............................27
11. Halodule wrightii distribution before (top) and after (bottom) Hurricane Irma ......................30
12. Syringodium filiforme distribution before (a) and after (b) Hurricane Irma ............................31
13. Proportion (%) of sites in each basin that had significant positive or negative change in
density ......................................................................................................................................32
14. Direction and intensity of change in total seagrass cover after Hurricane Irma. .....................33
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15. Thalassia testudinum cover before Hurricane Irma (top) and after Hurricane Irma
(bottom)....................................................................................................................................34
16. Halodule wrightii density before (top) and after (bottom) Hurricane Irma. ............................35
17. Syringodium filiforme density before (top) and after (bottom) Hurricane Irma. .....................36
18. Total Seagrass Cover by Braun-Blanquet value for before (top) and after (bottom)
Hurricane Irma .........................................................................................................................37
19. Aerial photography of the study area from one month after Hurricane Irma. Study basins
are outlined in yellow. The yellow star on the eastern edge of TKB marks the location of
sediment blow-outs. .................................................................................................................39
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CHAPTER 1
INTRODUCTION
Florida Bay is a saltwater portion of Everglades National Park positioned between the
southern river outlets, the Gulf of Mexico, and the Florida Keys (Figure 1). It supports a
tremendous variety of fauna, many of which are critical to the South Florida economy and
ecosystem. Not only is Florida Bay home to threatened and endangered species, including four
protected species of sea turtles and the endangered American crocodile (Crocodylus acutus) and
manatee (Trichechus manatus laterostris) (Florida Museum of Natural History), but it is also a
popular fishing destination that contributes over $800 million annually in economic activity and
nearly 9,000 jobs to South Florida (Fedler 2009). The sport fish behind this spending, as well as
the previously mentioned protected species, all depend, at least in part, on the seagrass
communities of Florida Bay for survival (Tilmant 1989; Fourqurean and Robblee 1999).
Hurricane Irma made US landfall through the Lower Florida Keys, with the eye of the
hurricane passing within 60 kilometers of the western extent of Florida Bay. Throughout the
southeastern US, the storm decimated crops and demolished coastal developments, leaving an
estimated $50 billion worth of damage in its wake (Cangialosi et al. 2018). The damage to land
structures and vegetation was obvious but what interested South Florida ecologists was more
difficult to observe; what effects did Hurricane Irma have on seagrass beds in Florida Bay?
The anthropogenically modified hydrology and polluted waters of Florida Bay are
severely limiting to the wetland and benthic communities within it. The past 100 years of coastal
development and hydrologic alteration to the Everglades have caused the “chronic state of low
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flow” in the northern and eastern sections of the bay that contribute to elevated salinity in the
estuarine basins (Brandt et al. 2016), while increased evaporation from rising sea surface
temperatures leads to a deficit of freshwater in the other subregions (Short and Neckles 1998;
Nuttle et al. 2000; Kelble et al. 2007). Prolonged periods of elevated salinity have been observed
to cause monocultures of Thalassia testudinum, which provide less suitable habitat for bay fish
species and are susceptible to widespread mortality (Thayer and Chester 1989; Bennet et al.
2005). The combination of high sea surface temperatures, hypoxia, and hypersalinity are
believed to be responsible for die-offs of T. testudinum like the one in 2015 (Borum et al. 2005;
Koch et al. 2007; Hall et al. 2016; National Park Service 2016).
In the months before Hurricane Irma struck in 2017, the basins that had been impacted by
the 2015 die-off still had water of higher salinity than average ocean water (South Florida Water
Management District 2018) and it was unclear whether the seagrass was recovering (Matt
Patterson, personal communication to the author, November 2017). Hurricanes are known to
reduce salinity in Florida Bay and are also expected to play a role in removal of sediment build-
up, increasing flow between basins (Zieman et al. 1989; Rudnick et al. 2005; Kelble et al. 2007).
It was predicted that total seagrass cover would increase after Irma and that community
composition would become more diverse as the influx in freshwater reduced bay salinity and
increased circulation, making conditions more suitable for the two target seagrass species
(Halodule wrightii and Syringodium filiforme) that are less salt-tolerant than T. testudinum.
Braun-Blanquet field survey methods were used to evaluate coverage and composition of
seagrass communities in two Florida Bay basins nearly a year after Hurricane Irma, at which
point colonization of scoured sediments could be seen. The basins used in this study had been
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previously assessed using these same field-survey methods to map the extent of the 2015 die-off,
the data from which was compared to the post-Irma survey to detect changes in seagrass status.
The purpose of this project is to record the changes in seagrass beds in Florida Bay after
Hurricane Irma. Submersed aquatic vegetation is considered a keystone species in Florida Bay
and, as such, is monitored by multiple agencies throughout South Florida (Brandt et al. 2016).
Visualization of the cover and community composition of seagrass beds in Florida Bay after
Hurricane Irma provide insight into the environmental conditions of the ecosystem and can be
used to track changes as conditions in the bay change, either as a result of restoration or of
climate change. Visualization of areas where there is no seagrass cover may also help guide
restoration projects in the future.
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Figure 1: Everglades National Park, Florida, USA. Park extent (green outline) and Florida
Bay extent (black dashed line) are shown.
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CHAPTER 2
BACKGROUND
2.1 Florida Bay Ecosystem
Resting at the southern outlet of the Everglades, Florida Bay is a subtropical body of
water that is known for its diverse plants and wildlife. The bay comprises nearly one third of
Everglades National Park and the Marjory Stoneman Douglas Wilderness, giving it the highest
level of protection assigned to US federal lands (National Park Service 2009). Florida Bay is a
popular destination for ecotourists while also serving as a place for both work and recreation for
locals. Sport fish such as tarpon (Megalops atlanticus), bonefish (Albula vupes), red drum
(Sciaenops ocellata), and spotted seatrout (Cynoscion nebulosus) live within the bay, making the
site popular for recreational fishing (Tabb and Manning 1961; Tilmant 1989; Powell 2003). In
addition, a rich community of piscivorous birds (e.g., wading birds, magnificent frigatebirds, and
osprey) and several protected vertebrate species, like American crocodiles (Crocodylus acutus)
and green sea turtles (Chelonia mydas), hunt within the bay or graze upon its seagrass beds
(Fourqurean and Robblee 1999). When seagrass beds significantly decrease in density, the bay
fauna decrease in abundance and diversity in response (Fourqurean and Robblee 1999).
Florida Bay is important to people from all walks of life, so the widespread impacts from
seagrass die-offs are an economic concern as much as an ecologic concern. Changes in climate
and Florida Bay hydrology throughout the 20th and 21st century have caused accelerated
evaporation and low circulation between basins, resulting in periods of hypersalinity, often at the
end of the dry season (November to April) (Fourqurean and Robblee 1999; Koch et al. 2007; Lee
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et al. 2016), the results of which have twice led to disastrous, widespread seagrass die-offs (Hall
et al. 2016). In basins with the lowest amounts of freshwater input, salinity has sometimes
reached 70 PSU, which is twice the average concentration of ocean water (Kelble et al. 2007).
While seagrasses are adapted to tolerate saline conditions, the mechanisms that control their
osmotic balance reduce photosynthetic efficiency (Touchette 2007) and increase their oxygen
demand for respiration (Koch et al. 2007).
The combination of elevated water temperature, decreased oxygen production from
photosynthesis, increased oxygen demand in plant tissues, and low circulation in the water
column creates hypoxic conditions, which encourages hydrogen sulfide production by anaerobic
microbes (Borum et al. 2005; Koch et al. 2007). Recent experiments have also found that, at
night, oxygen-deprived root tissue in Thalassia testudinum – the dominant seagrass species in
Florida Bay and the only species to survive long-term exposure to salinity over 40 practical
salinity units (PSU) – is especially susceptible to hydrogen sulfide toxicity via intrusion from
porewater when exposed to hypersaline conditions (Johnson et al. 2018). The decay of plant
tissues in anoxic conditions causes further hydrogen sulfide production, which results in a
positive feedback loop of decomposing seagrass leading to the mortality of the surrounding
seagrass (Carlson et al. 1994)
Hydrogen sulfide creates a “rotten egg” smell and yellow “fog” in the water (Hall et al.
2016; National Park Service 2016). The smell, along with floating masses of dead seagrass and
fish hurt the tourism industry and the decline in sport fish populations hurt the recreational
fishing industry. As stated by the Everglades Marine Team, “when Florida Bay is unhealthy, the
impacts are felt across South Florida” (National Park Service 2016).
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The literature suggests that precipitation from extreme storms can reduce hypersaline
conditions and remove sediment and detritus buildup in Florida Bay, relieving some of the
chemical stress on seagrass (Davis et al. 2004; Rudnick et al. 2005; Steward et al. 2006). This
would improve seagrass productivity, allow the less salt-tolerant seagrass species to populate the
seagrass communities, and provide higher-quality habitat for marine life (Thayer and Chester
1989; Bennett et al. 2005; Touchette 2007). However, these weather events also cause physical
stress on plants from wave activity and scouring and may reduce light availability to submerged
vegetation with sediment suspension and algal blooms from the released sediment nutrients
(Fonesca et al. 1998; Fourqurean et al. 1999; Davis et al. 2004; Rudnick et al. 2005; Steward et
al. 2006; Carlson et al. 2010; Cole et al. 2018).
2.2 Florida Bay Hydrology
Starting in 1881, the Florida Everglades underwent over 100 years of hydrologic
alterations in order to reclaim wetlands to accommodate the growing population of South Florida
(Light and Dineen 1994). The most intensive alterations, the Central & South Florida Project,
were completed in the 1950s-1970s; the project installed a network of canals, levees, pumps, and
water conservation areas with the goal of preventing floods and augmenting the local water
supply (Light and Dineen 1994; Ogden et al. 2005). Because of the hydrologic diversion, 6.4
billion kilograms of water bypass the Everglades, leaving the system with only 30% of its
original flow (USFWS 1999). The reduced flow is often credited for the high salinity and
increased salinity variability in northern sections of Florida Bay and influencing community
composition (Bennett et al. 2005; Rudnick et al. 2005).
While these developments allowed massive population growth and urbanization in South
Florida, by the early 1980s public concern for the ecosystem inspired changes in water
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management, starting with Florida Governor Graham’s launch of the “Save Our Everglades”
program in 1983 (Light and Dineen 1994). In 2000, the Comprehensive Everglades Restoration
Plan (CERP) set the guidelines for an interagency effort to restore the Everglades ecosystem
functions by imitating historic hydrology patterns (USACE and SFWMD 1999).
One of the many challenges of Everglades restoration is determining what the historic
hydrology patterns were; the quality and quantity of water in the freshwater Everglades and
Florida Bay were not well documented before the flow was diverted, so restoration efforts
depend upon statistical models and paleoecologic proxy data to estimate historic conditions
(Rudnick et al. 2005). The simulated historic conditions calculated by Rudnick et al. (2005) were
found to deliver 2.5-4 times as much freshwater to estuaries and wetlands as modern flow,
resulting in salinities in modern Florida Bay that are 5.3-20.1 PSU higher than before the
anthropogenic changes were made.
The complex network of mudbanks in Florida Bay restricts flow between basins, creating
a patchwork of variable water quality throughout the bay. Because of current hydrologic
conditions, only the northeastern section of the bay functions as a true estuary, where freshwater
from the Everglades mixes with saline water from the Gulf of Mexico; the rest of the bay
functions more like a tropical coastal lagoon, where most of the water flows in to the bay from
the Gulf of Mexico and precipitation is the primary source of freshwater (Rapozo 2001; Melo
and Lee 2012). Flow over shallow banks and through channels is primarily wind-driven and only
provide weak exchange between basins (Lee et al. 2006, 2008, 2016). As shown in Figure 2 (Lee
et al. 2016), basins in the western bay have greater exchange than interior and northeastern
basins, giving western basins a residence time of about 1 month compared to the 6-12-month
residence time in other sections (Lee et al. 2006, 2008, 2016). This restricted horizontal flow
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makes it unclear how restoration of freshwater inflow from the Everglades may affect the bay;
approximately 75% of all freshwater runoff from the Everglades flows into the northeast section
of the bay, where the low outflow stores it for an average residence time of one year (Lee et al.
2008). Nuttle et al. (2000) argue that, while increasing runoff from the Everglades would have
little effect on salinity in the central and west regions of Florida Bay – where seagrass die-offs
have occurred in the past – and would increase the variability of salinity in the southern section,
the effect of freshwater runoff in the bay could be greater if inflow is increased during periods
when evaporation exceeds precipitation. Similarly, Lee et al. (2006, 2016) suggest that
increasing flow to McCormick Creek during the dry season would reduce salinity in the north-
central subregion of the bay, possibly preventing hypersaline events.
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Figure 2: Annual mean volume transports (m3 s−1) for measured channels in Florida Bay (red
arrows and numbers), through the tidal passes between the Florida Keys (blue arrows and
numbers), and from mean river discharge (black arrows and numbers) (Lee et al. 2016).
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2.3 Plants Studied
The seagrass species most commonly
found in the Florida Bay are Thalassia
testudinum, Halodule wrightii and Syringodium
filiforme and they are the only species evaluated
in this research. Thalassia testudinum has wide,
long, green leaves which grow off an extensive
underground rhizome. Syringodium filiforme
produces cylindrical leaves that are similar in
length, if not longer and are a lighter shade of
green than that of T. testudinum. Halodule
wrightii produces the shortest leaves of the three,
which are flat and dark green to almost black in
color (Figure 3).
Thalassia testudinum is the most abundant
seagrass in Florida Bay and is considered the
dominant species in these communities (Fourqurean
et al. 2001). The size and low nutrient requirements
of this species allow it to out-compete the other
species in most conditions. It is also the most salt-
tolerant species investigated; studies have found
that the optimal salinity range of Thalassia
testudinum is likely between 20 and 40 PSU
Figure 3: Thalassia testudinum (top),
Halodule wrightii (middle), and
Syringodium filiforme (bottom).
Illustrations are by Phillips and Menez
(1988) and photographs are by Ken Dunton
(University of Texas).
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(Touchette 2007) and that in periods of prolonged salinity over 40 PSU, a monoculture of
Thalassia testudinum is observed (Bennett et al. 2005). Thalassia testudinum can be out
competed by S. filiforme in environments with low light availability or high phosphorous, which
is a limiting nutrient in Florida Bay and increases in concentration from east to west in this
system (Fourqurean et al. 1992). Halodule wrightii is the least sensitive to fluctuations in salinity
of the three species and is competitively excluded by the others in all environments expect
waters with high nutrient concentrations that experience widely fluctuating salinity or in the
wake of disturbance (Robblee et al. 1991).
While T. testudinum is dominant in succession, this species grows slowly (the median age
of single shoots of T. testudinum in Florida Bay is greater than 5 years) and typically reproduces
clonally, making it slow to repopulate areas defoliated by extreme weather (Robblee et al. 1991;
Peterson and Fourqurean 2001; Fourqurean et al. 2003). Halodule wrightii is considered a
pioneer species, which becomes established quickly in bare sediment if water quality is suitable
(Peterson and Fourqurean 2001). Considering the rates and requirements for growth of the three
species, it was expected that H. wrightii and S. filiforme, which could not tolerate the hypersaline
conditions present during the pre-Irma study, will exhibit new distributions after Irma. Halodule
wrightii was expected to be especially abundant in areas that were disturbed by the storm energy
where the dominant successional species – Thalassia testudinum – would not have had time to
reestablish after Hurricane Irma.
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2.4 Hurricane Effects
Hurricane Irma made U.S. landfall at Cudjoe Key as a category 4 hurricane on September
10, 2017. Florida Bay and Everglades National Park were located to the east of Irma’s path,
where the strongest winds of the storm were directed (Figure 4). While traveling past the western
Figure 4: Hurricane Irma's track in relation to Florida Bay and South Florida.
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edge of Florida Bay, the storm’s maximum recorded wind speed was 115 knots and minimum
pressure was 931 mb. Maximum inundation in the Lower Keys was nearly 2.5 m above the
normal tide line. Throughout the Keys, rainfall totals of 10-15 inches (25-38 cm) were reported
(Cangialosi et al. 2018).
In the United States, Irma caused 10 direct human deaths and 82 indirect human deaths.
The Middle and Lower Keys sustained the most damage, where an estimated 25% of all homes
were destroyed, while 90% of all homes were damaged. Irma cost an estimated $50 billion in
damages to structures and agriculture in the U.S. alone (Cangialosi et al. 2018).
Hurricanes and other extreme weather events are known to cause damage to natural and
man-made structures alike, but the short- and long-term effects of tropical cyclones (TCs) on
submerged aquatic vegetation can depend on the bathymetry, as well as the size, windspeed, and
traveling speed of the storm and water quality in the weeks after the storm passes (Craighead and
Gilbert 1962; Davis 2004; Carlson et al. 2010). It is uncommon for storm surge and shear
strength to damage fully submerged seagrasses, especially in a protected bay, but the sediments
disturbed by the storm can limit seagrass photosynthesis for several days and the resultant
deposition can bury leaves, either of which can cause seagrass mortality or reduced productivity
(Carlson et al. 2010). Because nutrients (especially phosphorous) accumulate in the carbonate
bay sediments, suspension of sediment also makes these nutrients available to algae, causing
blooms (Rudnick et al. 2005).
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In Florida Bay, some basins
are shallow enough that storm surge
and winds can scour seagrass beds,
removing sediment, seagrass root
networks, and detritus from the bay
floor (Davis et al. 2004). Wrack lines
– lines made of buoyant material that
clings to structures and is often used
to determine the peak water level of a
flood – that could be seen across
South Florida after Irma made it clear that seagrass had indeed been removed from the bay floor
by the storm (Figure 5).
While hurricanes cause devastation to structures and natural features, TCs are important
for regulating water quality and circulation in Florida Bay, with extreme weather events
providing a hydrologic pulse to the region (Davis et al. 2004; Rudnick et al. 2005; Steward et al.
2006). Buildup of carbonate sediment banks reduce flow between basins, contributing to the
hypersaline and hypoxic conditions during periods of high evaporation and low inflow to the
bay; storm surge is thought be able to remove enough sediment to restore flow while also
transporting excess, sediment-bound nutrients out of the bay (Rudnick et al. 2005). Freshwater in
Florida Bay is almost entirely acquired as rainfall, meaning that TC precipitation can also relieve
salt stress in confined basins (Nuttle et al. 2000). In 2018, the South Florida Water Management
District (SFWMD) reported salinities of 30-45 practical salinity units (PSU) throughout most of
the bay the month before Hurricane Irma, whereas one month after Irma most of the bay was
Figure 5: A wrack line composed entirely of dried seagrass,
mostly Thalassia testudinum, left by Hurricane Irma. This photo
was taken over 300 meters from Flamingo Beach, FL.
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recorded at 0-20 PSU (Figure 3). Five months after Irma, most of the bay still had salinity
concentrations below 30 PSU (SFWMD 2018), which is below the average salinity of ocean
water (35 PSU) and conducive to growth for all three of the target seagrass species (Fourqurean
et al. 2002; Touchette 2007).
Figure 6: Florida Bay salinity (PSU) one month before Hurricane Irma, one month after, and
five months after (SFWMD 2018).
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CHAPTER 3
MATERIALS AND METHODS
3.1 Study Sites
This study used two adjacent hydrogeologic basins – Rabbit Key Basin (RKB) and Twin
Key Basin (TKB) – that were impacted by the seagrass die-off in 2015 (McGinnis 2017). The
basins are located in the western subregion of Florida Bay (Figure 7), where precipitation is the
primary source of freshwater and most inflow comes from the Gulf of Mexico (Nuttle et al.
2000; Lee et al. 2016). Despite their close proximity, the basins differ substantially in water
quality and community compositions (Fourqurean et al. 2003), so all analysis of seagrass data in
this investigation will consider the basins individually.
Twin Key Basin was the first basin surveyed. The 65 km2 basin was turbid and affected
by abundant microalgae but the visibility was not impacted enough to prevent sampling. TKB
experiences hypersalinity more frequently than its western neighbor, RKB, due to its main
source of inflow coming from the bay’s most hypersaline basins (Lee et al. 2006, 2016). RKB is
smaller and shallower than TKB (47 km2) (Figure 8), but its position in the bay provides RKB
with direct water exchange with the Gulf of Mexico and the associated regulation of salinity (Lee
et al. 2016). RKB also had better visibility than TKB, most likely due to lower water column
nutrient concentrations limiting microalgal populations (Fourqurean et al. 1992). The southern
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edge of TKB had the greatest depth in the study area (nearly 3 meters below mean sea level) but,
overall, the majority of the study area is 1-2 meters deep (Figure 8).
Figure 7: Twin Key and Rabbit Key Basins and their position within Florida Bay.
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3.2 Sampling Method
In the pre-Irma, post die-off study, 35 study sites were randomly generated for each study
basin (McGinnis 2017). Of those 35 sites, 30 sites in TKB and 20 sites in RKB were surveyed
pre-Irma. For this Post-Irma study, researchers navigated to the same locations surveyed in the
pre-Irma study using a vessel-mounted GPS with waypoints uploaded using the DNRGPS
application by the Minnesota Department of Natural Resources. Of the sites that had Pre-Irma
data, all but two sites were surveyed – two sites in RKB were in water too shallow for the
research vessel that was larger than the one used for the Pre-Irma survey – resulting in a total of
48 study sites (Figure 9).
Figure 8: Bathymetry of Rabbit Key and Twin Key Basins. Contour interval: 0.25 m. Digital
Elevation Model provided by National Centers for Environmental Information (NCEI).
Vertical error estimated by NCEI to be no more than 15 cm.
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For each study site, once the vessel reached the waypoint, a team member dropped a soft-
weighted buoy to mark the spot. A pair of snorkelers then attached two 30-meter transect tapes to
the buoy and swam in different, random directions. Each diver haphazardly dropped a 0.25 m²
PVC quadrat at 5-meter intervals (5, 10, 15, 20, and 25 meters from the buoy) and analyzed the
seagrass cover within the quadrat where it landed.
Density was assessed using the Braun-Blanquet cover-abundance method, which ranks
shoot density on an ordinal scale; a 5 on the scale represents 75-100% cover, 4 represents 50-
75%, 3 represents 25-50%, 2 represents 10-25%, and 1 represents less than 10%. If seagrass is
found in the transect but there is too little for it to provide cover, a ranking of 0.5 is used to
Figure 9: Location of sampling sites in RKB and TKB.
21
indicate that that there were several shoots of seagrass or 0.1 indicates that there was one shoot
of seagrass. Each new snorkeler was trained by an experienced team member to ensure that
Braun-Blanquet values assigned by different researchers corresponded to the same approximate
density throughout the study. Snorkelers recorded the Braun-Blanquet value describing cover-
abundance of the three target-species at each of the 10 transects upon their return to the vessel.
The Braun-Blanquet method is used in this study for two reasons: the pre-Irma survey
utilized this method and changing the metric would make the validity of comparisons between
datasets questionable (Fonesca et al. 1998), and because it is a rapid, easily used method which is
invaluable when the funding, resources, and weather limits the number of days that can be spent
collecting data. If a more time-intensive metric, such as shoot density, had been found for each
site, a much smaller number of sites would have been used in this study and, as a result,
interpolation would have been less robust. This survey method is also preferred by National
Parks officials over more destructive methods, such as measuring biomass or collecting dredge
samples, both of which kill the seagrass sampled (Matt Patterson, personal communication to
author, November 2017). In comparison, Braun-Blanquet surveying is minimally destructive,
with the anchor of the boat inflicting the only obvious damage to seagrass. Other less direct
methods of quantifying seagrass distribution involve remote sensing, which was not appropriate
for this research because the type of benthic vegetation visible in remotely sensed imagery is
nearly impossible to determine for Florida Bay. Not only is the water often too turbid to see the
bottom, but several macroalgae species can be seen throughout the study area, some of which
drift freely along the sea floor, making it possible for images taken hours apart to have different
distributions of submersed aquatic vegetation (Matt Patterson, personal communication to
author, November 2017). The community structure of seagrass also limits the utility of remote
22
sensing in this environment, as seagrass can grow in multiple stories and all three species can be
present at a site in a nearly-even distribution (Fourqurean et al. 2002); spectral signatures could
be used to identify species in areas with more homogenous composition but, for heterogenous
communities like those in the western Florida Bay, it would not have been useful. In the context
of this research, the only practical survey method was Braun-Blanquet.
3.3 Spatial Analysis
Change analysis for each species was performed in IBM® SPSS®. While the Braun-
Blanquet cover-abundance method is useful for rapidly approximating density in the field, the
ordinal nature of the data requires use of non-parametric tests for determining significant change.
The Kruskal-Wallis test was chosen to determine which sites had significant (P=0.05) difference
in species density and total density – the sum of the three species densities in each sample –
between the two study periods. The test was performed for all four density variables for each of
the 48 sites (192 Kruskal-Wallis calculations total). Tests used the 20 sample values describing
each density variable – ten samples from the pre-Irma survey and ten samples from the post-Irma
survey – a total of 3,840 data points. The recorded Kruskal-Wallis test statistics indicated
whether a variable had the same mean rank across the two study periods; test statistics of P<0.05
meant that the data distribution between the two time periods were significantly different.
Sites that had significant change in total density were plotted in ArcMap. Maps indicating
zones of significant change in density were created using only these sites; the difference in
medians from before and after Hurricane Irma was used to indicate if the change was positive or
negative and inverse Distance Weighting (IDW) was used to create a raster for these points.
Density of seagrass and density of individual species was mapped by plotting the median
Braun-Blanquet score for the density variable at the coordinates assigned to each site. A raster
23
surface was created for each species by basin using Inverse Distance Weighting (IDW)
interpolation with a cell size of 45 meters. This cell size was selected because it produced cells of
the same area as a circle with a radius of 25 meters – the furthest sampling distance from the
buoy at each site. Other IDW settings were kept at default, with a distance power of 2 and a 12-
point search parameter. The rasters were processed to the extent of each basin using appropriate
polygons. After extraction by a mask of the same basin polygons, the raster data was classified
by Braun-Blanquet value. Maps are shown with an overlay outlining locations of shallow banks,
where intertidal communities are likely to differ from benthic seagrass meadows and where
sampling was sometimes impossible in the 25-foot NPS research vessel. The shallow banks
polygon layer is provided by the Florida Fish and Wildlife Conservation Commission-Fish and
Wildlife Research Institute.
The validity of statistical relationships between points from ordinal data is a topic of
debate in statistics (Allen and Seaman 2007). The ranges of percent cover that each Braun-
Blanquet class represents are not all equal and the distribution of the data is not normal, leading
to my assertion that it is not appropriate to derive statistical relationships from this data as if it
were interval data. For these reasons, Kriging – an interpolation method that uses the variance of
the data to create maps that are weighted by the statistical significance of data over distance –
was not used, despite its reputation as a more robust interpolation method. Because of the ordinal
nature of the Braun-Blanquet values, spatial interpolation should only be used to determine the
boundaries between the classes; the area between boundaries cannot be interpolated to non-
integer numbers (Feoli and Orloci 2012). While IDW interpolation may oversimplify the
geospatial patterns present, it produces a visualization of general trends in the study area that
may be useful for natural resource management.
24
CHAPTER 4
RESULTS
After Hurricane Irma, overall density of seagrass in the study area became more
contiguous, with fewer patches of bare sediment (no seagrass) and sparse cover (<10% seagrass
cover) observed (Figure 10). Distribution of the three seagrass species increased in both basins in
all but one variable; S. filiforme in Twin Key Basin (Table 1). The pioneer species Halodule
wrightii experienced the greatest change in distribution, as it was found in 25% more samples in
RKB and 6.3% more samples in TKB after Irma. Thalassia testudinum grew in the greatest
densities, providing dense cover (>50% seagrass cover) at 40.6% of samples in RKB and 18.3%
of samples in TKB.
Twin Key Basin was overwhelmingly dominated by T. testudinum (Table 1). Only 0.3%
of samples in this basin reported more than 10% cover from any other species, which in this case
was Halodule wrightii. While density of H. wrightii was low in TKB, its distribution after Irma
spanned most of the basin (Figure 11). Syringodium filiforme, however, was almost completely
absent in TKB, where only one pre-Irma sample at one site reported its presence. With a Braun-
Blanquet value of 0.1, the single shoot of S. filiforme found in TKB was not enough to change
the site’s median S. filiforme density, so no map of S. filiforme density in TKB was produced.
T. testudinum was also the most abundant seagrass species in RKB, but H. wrightii and S.
filiforme appeared concurrently in greater densities in this basin than in TKB (Table 1). Both H.
wrightii and S. filiforme were distributed across the basin, with few patches where they were
absent (Figures 11, 12).
25
Kruskal-Wallis tests determined significant change in density of seagrasses in both
basins. The difference between post-Irma and pre-Irma median densities for each site provided
information on the direction of change at these significant sites (Figures 13, 14). While positive
and negative changes were seen throughout the study area, Halodule wrightii was the only
species to not experience significant negative change after Hurricane Irma. Syringodium filiforme
experienced equal proportions of positive and negative change while T. testudinum and total
seagrass density in both basins had at least 10% more sites experience positive change than
negative change.
Thalassia testudinum density became more uniform throughout the study area, as areas of
sparse cover and areas of dense cover both approached more moderate densities (Figure 15).
Density of H. wrightii increased in both basins, though it provided no more than 10% cover in
most of the study area (Figure 16). Basin-wide density of S. filiforme did not change noticeably,
though the patterns of density shifted after Hurricane Irma (Figure 17). Changes in individual
species’ density influenced the total seagrass cover, which became more uniform after Irma
(Figures 10, 18).
The accuracy of each density interpolation, calculated by Root Mean Squared Error
(RMSE), is presented in the Appendix. Error was measured using the difference between
interpolated values and a set of observations not used to create the interpolated maps;
approximately 10% of the sites from each basin were reserved for use in the RMSE. The most
accurate interpolation was the post-Irma map of total cover in RKB (0.035 RMSE) and the least
accurate interpolation was the pre-Irma map of total cover in TKB (1.517 RMSE). Because of
the “patchy” nature of seagrass beds in the study area, this amount of error is expected; the
interpolation maps represent the median density of each species, but bare patches and unusually
26
dense patches should be anticipated (Pramova et al. 2013). More reliable maps, made by
classifying total cover into the broader categories of “Bare,” representing no cover, “Sparse,”
representing less than 10% cover, “Moderate,” representing 10-50% cover, and “Dense,”
representing more than 50% cover, are shown for before and after Irma (Figure 10).
27
Figure 10: Total seagrass cover defined by broad cover categories. Bare sediment is 0% cover;
Sparse cover is <10%; Moderate cover is 10-50%; dense cover is >50%.
28
Table 3(a-b): Braun-Blanquet assessment of seagrass density (D) for all samples, separated by species and basin. 1a contains pre-
Irma data and 1b contains post-Irma. Adapted from Fourqurean et al. (2001).
29
30
Figure 11: Halodule wrightii distribution before
(top) and after (bottom) Hurricane Irma.
31
Figure 12(a-b): Syringodium filiforme distribution before (a) and after (b)
Hurricane Irma.
32
Figure 13: Proportion (%) of sites in each basin that had significant positive or negative
change in density.
33
Figure 14: Direction and intensity of change in total seagrass cover after Hurricane
Irma.
34
Figure 15: Thalassia testudinum cover before Hurricane Irma
(top) and after Hurricane Irma (bottom).
35
Figure 16: Halodule wrightii density before (top) and after (bottom) Hurricane Irma.
36
Figure 17: Syringodium filiforme density before (top) and after (bottom)
Hurricane Irma.
37
Figure 18: Total Seagrass Cover by Braun-Blanquet value for before
(top) and after (bottom) Hurricane Irma.
38
CHAPTER 5
DISCUSSION
Hurricane Irma produced strong western winds over Florida Bay, which directed
powerful waves towards the northwest within both basins that may have caused the decrease in
seagrass density visible in the west and northwest sections of both basins (Figure 13). In TKB,
T. testudinum cover increased after Hurricane Irma in the north-northeast portion of the basin.
This region was severely impacted by the seagrass die-off in 2015, so positive change in cover
indicates that conditions became more hospitable for T. testudinum in this part of the bay (Matt
Patterson, personal communication to author, November 2018). The northern section of TKB
receives flow from Whipray Basin and other north-central basins of that experience some of the
most extreme, chronic hypersalinity in Florida Bay (Lee et al. 2006, 2016), so regulation of
salinity and improved flow due to the influx of freshwater from Hurricane Irma may have had
greater impact in this area than in other parts of the study area.
Other changes in cover may be explained by the bathymetry of these basins. RKB is
confined from the southwest by Ninemile Bank, which is believed to have shielded much of the
basin from the most intense wave action. Conversely, the eastern inlets to TKB are less protected
by banks and are shallow, leaving this region exposed to scouring waves. As seen in Figure 17,
this region did not have uniform dense cover before Irma, meaning that existing sediment could
have been removed or suspended by wave action. As a result, sediment experienced a “blow-out”
at this location, possibly taking the seagrass with it; aerial photography from one month after
Irma shows plumes of suspended sediment going from the banks and narrow channels into TKB
39
(Figure 18). If physical removal by wave energy did not cause the decline in T. testudinum
density on the eastern edge of TKB, then the reduced light penetration from suspended sediment
and algal blooms is another possible cause (Short and Neckles 1999; Rudnick et al. 2005). This
area is also noteworthy because, while T. testudinum density significantly decreased, H. wrightii
has increased in density there after Irma (Figure 15). The transition from T. testudinum to H.
wrightii aligns with the literature, which identifies H. wrightii as a successful pioneer species
after disturbance (Peterson and Fourqurean 2001). This also indicates that water quality has
changed to be more suitable to H. wrightii since the pre-Irma survey; H. wrightii has been found
to decline in water with salinity over 30 PSU and to be absent over 40 PSU (Bennett et al. 2005;
Touchette 2007).
Figure 19: Aerial photography of the study area from one month after Hurricane Irma. Study
basins are outlined in yellow. The yellow star on the eastern edge of TKB marks the location of
sediment blow-outs.
40
The distribution of H. wrightii and S. filiforme also changed, with H. wrightii (Figure 8)
becoming denser throughout the center and southeastern portion of the basin and S. filiforme
(Figure 9) becoming denser in the northern portion of the basin. These changes in density could
have been caused by improved nutrient or light availability, which is evidenced by the inverse
relationship between T. testudinum and S. filiforme in this basin; S. filiforme has been found to
out-compete T. testudinum in waters with low light or high phosphorous concentrations, which is
the limiting nutrient for primary production in the western section of the bay (Fourqurean et al.
1992).
The change in median total density of both basins (Figure 10), makes it clear that after
Hurricane Irma, both basins showed less area that had bare sediment or sparse cover (less than
10% cover, or 0-1 in Braun-Blanquet values, as mapped). The post-Irma map also shows more
uniform coverage, indicating that cover is more contiguous after Irma. More contiguous, dense
cover can stabilize sediments more efficiently, preventing the resuspension of sediment and
nutrients, which lead to algae blooms (Rudnick et al. 2005; Hansen and Reidenbach 2017).
The data shows that there are sections of the bay that experienced a significant loss of
seagrass but there are other areas that not only experienced a significant increase in seagrass
density, but also demonstrated higher species richness when water quality became more
hospitable to H. wrightii and S. filiforme, which forms more suitable habitat for fish and other
bay fauna (Bennett et al. 2005). This research contributes new data to the literature on Florida
Bay ecology by providing evidence that tropical cyclones may help accelerate recovery after
seagrass die-off events. Factors that may have influenced seagrass recovery include 1) dilution of
hypersaline water by precipitation and storm surge and 2) removal of built-up sediments from
banks and channels, allowing more water exchange between basins.
41
CHAPTER 6
CONCLUSION
The precipitation-dependency and low flow between basins in Florida Bay makes the
region prone to water deficit, leading to variable salinity. In cases of prolonged hypersalinity,
interaction between environmental stressors can cause wide-spread seagrass die-offs (Borum et
al. 2005; Koch et al. 2007; Johnson et al. 2018). After an extensive die-off event in 2015,
hypersaline water that was high in hydrogen sulfide concentration and low in dissolved oxygen
due to the decomposing seagrass remained trapped in the confined basins of the bay. When
Hurricane Irma passed Florida Bay in 2017, scientists from multiple agencies in South Florida
were interested to see the positive and negative effects that the event might have on the
hydrologically challenged system.
While the storm caused decline in seagrass density in some parts of the study area,
seagrass cover in the majority of the study area became more uniform and more diverse. This
research shows that, throughout the study basins, the overall seagrass density increased and there
was less area with bare sediment. I also found that the non-dominant species Halodule wrightii
and Syringodium filiforme began growing in greater densities in some areas where they had
already been established and had colonized some patches of sediment that had been left bare
after Hurricane Irma. There are still portions of both basins that show a significant reduction in
seagrass density after Irma but the positive influence the storm had on seagrass communities
after a die-off event is an ecological silver lining to this devastating storm.
42
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47
APPENDIX
Accuracy Assessment
The accuracy of each interpolated map was performed using 10% of the data points, which were
reserved during interpolation. Root Mean Squared Error of each site is reported in Table 3. Error
was calculated as the difference between observed values and the nearest boundary of the range
used to define the predicted values.
Table 4: Root Mean Square Error for each map by basin (when applicable).
RABBIT KEY BASIN TWIN KEY BASIN
T. TESTUDINUM pre-Irma 1.051855741 1.041633333
T. TESTUDINUM post-Irma 0.637965908 0.777281588
H. WRIGHTII pre-Irma 0.360555128 N/A
H. WRIGHTII post-Irma 1.457909634 0.057735027
S. FILIFORME pre-Irma 0.070710678 N/A
S. FILIFORME post-Irma 0.320156212 N/A
TOTAL COVER pre-Irma 0.285745516 1.517124473
TOTAL COVER post-Irma 0.035355339 0.777281588