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Importance of Storm Events in Controlling EcosystemStructure and Function in a Florida Gulf Coast EstuaryStephen E. Davis, III†*, Jaye E. Cable‡, Daniel L. Childers§, Carlos Coronado-Molina††, John W. Day, Jr‡,Clinton D. Hittle‡‡, Christopher J. Madden††, Enrique Reyes§§, David Rudnick††, Fred Sklar4

†Department of Wildlife &Fisheries Sciences

Texas A&M UniversityCollege Station, TX 77843-

2258sedavis@tamu.edu

‡‡Department ofOceanography and CoastalSciences

Coastal Ecology InstituteLouisiana State UniversityBaton Rouge, Louisiana

70803jcable@lsu.edu,

johnday@lsu.edu

§Department of BiologicalSciences & SERC

Florida InternationalUniversity

Miami, Florida 33199childers@fiu.edu

††Everglades SystemResearch

South Florida WaterManagement District

3301 Gun Club RoadWest Palm Beach, Florida

33416cmadden@sfwmd.gov,

drudnic@sfwmd.gov,fsklar@sfwmd.gov

‡‡United States GeologicalSurvey

Center for Water andRestoration Studies

9100 NW 36th St. Suite # 107Miami, Florida 33178cdhittle@usgs.gov

§§Department of Geology andGeophysics

University of New Orleans2000 Lakeshore Dr.New Orleans, LA 70148ereyes@uno.edu

ABSTRACT

DAVIS, S.E., III; CABLE, J.E.; CHILDERS, D.L.; CORONADO-MOLINA, C.; DAY, J.W., JR.; HITTLE, C.D.; MAD-DEN, C.J.; REYES, E.; RUDNICK, D., and SKLAR, F., 2004. Importance of storm events in controlling ecosystemstructure and function in a Florida gulf coast estuary. Journal of Coastal Research, 20(3), 000–000. West Palm Beach(Florida), ISSN 0749-0208.

From 8/95 to 2/01, we investigated the ecological effects of intra- and inter-annual variability in freshwater flowthrough Taylor Creek in southeastern Everglades National Park. Continuous monitoring and intensive samplingstudies overlapped with an array of pulsed weather events that impacted physical, chemical, and biological attributesof this region. We quantified the effects of three events representing a range of characteristics (duration, amount ofprecipitation, storm intensity, wind direction) on the hydraulic connectivity, nutrient and sediment dynamics, andvegetation structure of the SE Everglades estuarine ecotone. These events included a strong winter storm in November1996, Tropical Storm Harvey in September 1999, and Hurricane Irene in October 1999. Continuous hydrologic anddaily water sample data were used to examine the effects of these events on the physical forcing and quality of waterin Taylor Creek. A high resolution, flow-through sampling and mapping approach was used to characterize waterquality in the adjacent bay. To understand the effects of these events on vegetation communities, we measuredmangrove litter production and estimated seagrass cover in the bay at monthly intervals. We also quantified sedimentdeposition associated with Hurricane Irene’s flood surge along the Buttonwood Ridge. These three events resulted indramatic changes in surface water movement and chemistry in Taylor Creek and adjacent regions of Florida Bay aswell as increased mangrove litterfall and flood surge scouring of seagrass beds. Up to 5 cm of bay-derived mud wasdeposited along the ridge adjacent to the creek in this single pulsed event. These short-term events can account fora substantial proportion of the annual flux of freshwater and materials between the mangrove zone and Florida Bay.Our findings shed light on the capacity of these storm events, especially when in succession, to have far reaching andlong lasting effects on coastal ecosystems such as the estuarine ecotone of the SE Everglades.

INTRODUCTION

Pulsed events such as winter storms, tropical storms, andhurricanes affect multiple structural and functional attri-butes of coastal ecosystems including hydrodynamics (PEREZ

03-0072R received 02 July 2003; accepted in revision 19 September2003.

* Corresponding author.

et al. 2000; PITTS 2001; VALLE-LEVINSON et al. 2002), wa-ter quality (RYBCZYK et al. 1995; FOGEL et al. 1999), emer-gent vegetation cover (DOYLE et al. 1995; RYBCZYK et al.1995), nutrient fluxes (RYBCZYK et al. 1995; PAERL et al.2001), sediment dynamics (CHILDERS and DAY 1990; CA-HOON et al. 1995; PEREZ et al. 2000), and biotic communitystructure (MICHENER et al. 1997; CABELLO-PASINI et al.2002). However, the importance of such events in regulating

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estuarine function over long time scales is not yet fully un-derstood (DAY et al., 1997). This is partly attributable to thepaucity of information on the response of a given coastal eco-system to a range of these events.

Between 1995 and 2001, we studied the ecological effectsof intra- and inter-annual variability in freshwater flow alongthe estuarine ecotone of the SE Everglades. This region ischaracterized by numerous mangrove creeks and shallowponds linking the freshwater Everglades marshes to FloridaBay (Figure 1). Over the years, continuous monitoring andintensive sampling studies overlapped with an array ofpulsed events that resulted in immediate and sustained im-pacts on the physical, chemical, and biological attributes ofthis region. Here, we provide a quantitative analysis of mul-tiple pulsed events, representing a range of storm traits (du-ration, amount of precipitation, intensity, wind direction). Weexamined how these events: 1) regulated the hydraulic con-nectivity between the mangrove ecotone and NE Florida Bay:2) controlled materials exchange at the mangrove–bay inter-face, and: 3) shaped vegetation dynamics throughout the SEEverglades.

The SE Everglades Mangrove System

Taylor Creek drains much of Taylor Slough in SE Ever-glades National Park (Figure 1) and has been the focus ofecological research and monitoring over much of the past 7years (RUDNICK et al., 1999; SUTULA et al. 2001; DAVIS etal. 2001a; 2001b; CORONADO-MOLINA et al., 2003; DAVISet al. 2003; SUTULA et al. 2003). This relatively short (, 5km in length) and narrow (ranging from 5–15 m), creek linksseveral small, shallow ponds from lower Taylor Slough to Lit-tle Madeira Bay and Florida Bay. As with many other creeksthat drain this region, Taylor Creek cuts through an em-bankment known as the Buttonwood Ridge before emptyinginto the bay.

Annual hydrographic variation in lower Taylor Creek is afunction of wind-driven flow and freshwater drainage fromTaylor Slough and the C-111 Canal system to the north (SU-TULA et al. 2001). Within Taylor Creek and the estuarineecotone, tidal ranges are less than 0.1 m, and are frequentlymasked by wind forcing. Southerly winds can push high sa-linity bay water more than 1 km into the mangrove fringewetlands, while northerly winds push freshwater out into Lit-tle Madeira and Florida Bays. Wet seasons (June–November)are usually characterized by high, positive freshwater dis-charge, which can also negate tidal exchange. Dry seasons(December–May) are characterized by poly- to euhaline con-ditions and low net discharge from Taylor Creek (Figure 2).In very dry years, this seasonality can result in an annualrange of salinity as high as 52 PSU in Little Madeira Bay,representing the highest annual range in eastern Florida Bay(ROBBLEE et al., 1989).

At the northern end of the estuarine ecotone, the wetlandvegetation grades from freshwater macrophyte species, such assawgrass (Cladium jamaicense) and spikerush (Eleocharis spp.),into a near monospecific stand of dwarf red mangrove (Rhizo-phora mangle). This dwarf mangrove environment is wide-spread throughout the SE Everglades estuarine ecotone and is

characterized by mature trees less than 2 m in height. Thefringe-form of R. mangle borders the lower reach of TaylorCreek. Immediately behind this zone and along the ButtonwoodRidge there is a basin mangrove forest dominated by button-wood (Conocarpus erectus) and white and black mangroves (La-guncularia racemosa and Avicennia germinans).

Regional Climate and Storm History

South Florida has a humid, subtropical climate influenced byprevailing northeast trade winds and the adjacent marine sys-tems of the Gulf of Mexico, Florida Bay, and the Atlantic Ocean.The region experiences a dry season typically from Novemberto May and a wet season nominally from May through October.Superimposed on these two seasons are the frontal season (dur-ing the early dry season) and the hurricane season (during thewet season). Nearly every year, south Florida experiences atleast one tropical storm and regular winter frontal passages(NEUMANN et al., 1978; SIMPSON and RIEHL, 1981; WAN-LESS et al., 1994). The recurrence interval of these events alonesuggests their importance in shaping the coastal Everglades,and mounting data support this notion (EGLER, 1952; ROB-BLEE et al., 1991; MCIVOR et al., 1994; LORENZ et al., 2001).Studies in other coastal regions of the world also indicate theimportance of these types of storm events in driving coastal eco-system structure and function (HENSEL et al., 1998; FOGEL etal., 1999; PEREZ et al., 2000; SHIAH et al., 2000; CABELLO-PASINI et al., 2002).

Of the worst storms to make landfall in the U.S. during the20th Century, three greatly impacted south Florida both eco-nomically and ecologically: 1) Labor Day Hurricane in 1935(Florida Keys), 2) Hurricane Donna in 1960 (Florida Keysand Ft. Meyers), and 3) Hurricane Andrew in 1992 (Miami-Dade County). The frequency of a direct hit by hurricanes ofthis magnitude in the Florida Keys and Florida Bay is ap-proximately once every 25–30 years (GENTRY, 1974). Duringthe 6-year period of our study (1995 to 2001), south Floridaeither was hit directly or was affected by at least eight ofthese tropical events (Table 1). In this synthesis we highlightthe effects of three of these storms.

1996 Winter Storm (wind event)

A lengthy winter storm, from 9–19 November 1996, wascharacterized by persistent gale force winds from the north-east blowing at about 15–20 m sec21. Negligible precipitationwas associated with this storm, but sustained winds forcedwater out from the freshwater Everglades, through TaylorCreek, and into Florida Bay for ten consecutive days. Whenthe storm passed, the winds reversed direction and water re-entered Taylor Creek from Little Madeira Bay. We use thisevent to demonstrate the effects of intense, long durationwind events with little precipitation.

Tropical Storm Harvey

Harvey formed shortly after Hurricane Floyd (19 Septem-ber 1999) in the eastern Gulf of Mexico and moved towardwest central Florida in the days following. As it approachedthe Florida peninsula it turned southeast and traversed the

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Figure 1. Map of south Florida highlighting the different zones (freshwater, mangrove, and Florida Bay) of Everglades National Park. Lower imagedepicts the mangrove zone of lower Taylor Slough and the embankment (i.e. Buttonwood Ridge) that borders Little Madeira Bay. Sites at the upperreach (TS/Ph 6) and the mouth of Taylor Creek (TS/Ph 7) are indicated.

Everglades (21 September) with the characteristics of a fron-tal wave. Wind velocities were generally less than 10 m sec21

and maximum precipitation (25 cm) was reported in Naples,FL. We observed a total of about 5 cm precipitation near oursite during the storm’s passage (Figure 2b). We use this eventto demonstrate the effects of short duration wind events withintermediate precipitation.

Hurricane IreneAfter moving across western Cuba as a tropical storm,

Irene strengthened into a hurricane in the Florida Straits on

15 October 1999. As a Category 1 hurricane (Saffir-SimpsonScale), Irene hit the lower Florida Keys at 09:00 15 October,then crossed Florida Bay and made landfall again near CapeSable, Florida at 16:00 that same day. The storm path con-tinued across Miami-Dade and Broward Counties and exitedthe Florida peninsula on the Atlantic coast near Jupiter, FL,on 16 October 1999. Hurricane Irene brought substantial pre-cipitation (as much as 44 cm measured in Boynton Beach,FL) and tropical storm force winds (, 27 m sec21) to southFlorida. We observed about 11 cm total precipitation nearTaylor Creek from H. Irene (Figure 2b). We use this event to

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Figure 2. Atmospheric conditions and Taylor Creek stage, discharge, and salinity during periods from October 1996–January 1997 and August 1999–December 1999. In the second set of graphs, barometric pressure is plotted with a dashed line and air temperature with a solid line. In the bottom setof graphs, salinity is plotted with a gray line and water temperature with a black line. (Left Panel): During November 1996, a winter wind storm pushedwater out of Taylor Creek 10 days before a wind shift reversed flow and water surged into the mangrove fringe. (Right Panel): Hurricane Floyd did notcross the Everglades, but its outer rain bands crossed south Florida in mid-September. Tropical Storm Harvey represented a relatively brief precipitationand runoff event in the Everglades 1.5 weeks later. One month later, Hurricane Irene passed over the SW Everglades. Stage values in Taylor Creek rosesharply from about 0.5 m to an annual maximum of 7.4 m within 12 hours.

Table 1. Caption?

Year

# ofNamed

Storms**

# MakingLandfall

in Florida

Named StormsAffecting

South Florida Comments

1995 19(11)

4 H. ErinT. S. Jerry

Second busiest Atlantic hurricane season since1871; H. Allison and Opal hit Panhandle;Erin and Jerry dropped rainfall in the areanorth of the Everglades National Park(ENP).

1996 13(9)

1 none Above average hurricane season; H. Josephinehit Panhandle

1997 7(3)

0 none Lackluster hurricane season, H. Erika broughtsome rainfall to north Florida

1998 14(10)

3 H. GeorgesH. Mitch

H. Earl struck north Florida; Mitch crossednorth of the ENP; Georges struck the lowerFlorida Keys

1999 12(8)

2 H. Floyd, T. S.Harvey, H.Irene

H. Floyd did not make landfall in Florida, butrain bands extended over S. Florida

2000 15(8)

3 none H. Gordon, T. S. Helene and Leslie hit NorthFlorida

2001 15(9)

1 H. Gabrielle Busy season overall, though most storms didnot affect Florida

* Data compiled from National Climatic Data Center (http://www.ncdc.gov) for Atlantic hurricane tracks since 1886.** Number in parentheses indicates storms of hurricane strength.

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demonstrate the effects of intense, short duration windevents with high precipitation.

METHODS

Hydrologic and Atmospheric Data

Stage, discharge, salinity, and water temperature werecontinuously measured at two U.S. Geological Survey (USGS)monitoring stations along Taylor Creek, one at the mouthand the other approximately 2.5 km upstream near the in-terface with the freshwater Everglades. We used the stationat the mouth of Taylor Creek (Figure 1) to document hydro-logic exchange between the mangrove ecotone along TaylorCreek and northeast Florida Bay during each of these ob-served storm events and in relation to annual measurements.Measurements of water velocity, stage, salinity, and watertemperature were taken every fifteen minutes and recordedby a data logger. These data were subsequently transmittedvia satellite to the USGS office in Miami, FL. Water velocitywas measured with an acoustic velocity meter (AVM), stagewas measured within a stilling well, and salinity and tem-perature were measured near the surface and bottom of thecreek (HITTLE et al. 2001). All stage measurements are re-ported relative to the North American Vertical Datum of1988 (NAVD ‘88).

We obtained wind speed and direction, precipitation, airtemperature, and barometric pressure from an atmosphericweather station located in Joe Bay (northeast of Little Ma-deira Bay). This station collected data for the National ParkService and the National Oceanographic and AtmosphericAdministration at 15-minute intervals. Wind data were vec-tor averaged to obtain daily values. These meteorologicaldata were used to document the effects of each storm and arepresented with the hydrologic data to present a completephysical picture of the wetlands-bay coupling before, during,and after these events (Figure 2).

Nutrient Sampling and Analyses

Since April 1996, daily water samples (DWS) have beencollected at the mouth of Taylor Creek (see TS/Ph-7, Figure1) to track temporal changes in salinity, total nitrogen, andtotal phosphorus in this system. Surface water samples werecollected by an automated sampler and retrieved at intervalsof 3 to 4 weeks. Until 30 September 1998, the samplingscheme produced a daily, 1-L water sample that integratedfour 250-mL sub-samples taken every six hours. Beginning 1October 1998, a tri-daily sampling program was implementedin which the 250-mL sub-samples were collected every 18hours representing the dawn (0600), dusk (1800), noon, andmidnight phase of each three-day cycle. Quarterly, from Au-gust 1996 to June 2001, we also performed intensive watersamplings (IWS) in which duplicate, 1-L surface water sam-ples were collected at six-hour intervals for a period of tendays.

In the DWS program, we analyzed samples for total nitro-gen (TN) and total phosphorus (TP). In the IWS program, wefiltered (Whatman GF/F) a portion of each sample and ana-lyzed for soluble reactive phosphorus (SRP), ammonium

(NH41), nitrate 1 nitrite (NOx

2), and dissolved organic car-bon (DOC). Unfiltered IWS samples were analyzed for TN,TP, and total organic carbon (TOC). See DAVIS et al. (2001a)for description of the analytical methods used.

For both sampling schemes (DWS and IWS), nutrient ex-change between the estuarine ecotone and the bay was cal-culated using concentration and discharge data from themouth of Taylor Creek. Daily fluxes of nutrients were quan-tified as the product of mean daily discharge and daily nu-trient concentration and integrated over an entire year toobtain a total annual flux.

Flow-Thru Sampling and Mapping

Spatial patterns in the Taylor Creek–Little Madeira Baysystem were measured periodically with a high-speed ‘‘data-flow’’ measurement apparatus developed for mapping physi-co-chemical parameters in shallow aquatic systems (MAD-DEN and DAY 1992). This integrated instrument system wasmounted to a boat and used to sample water temperature,conductivity, salinity, and water clarity (beam transmittance)while running tightly gridded transects throughout LittleMadeira Bay and the lower Taylor Creek on October 20, 1999(5 days after Hurricane Irene). The same transects were runone year later on October 19, 2000, after a period of nostorms, to provide a reference dataset for comparison. In bothsurveys, samples were taken at 4-sec intervals (approximate-ly every 2 to 8 m, depending on boat speed). An integratedglobal positioning system (GPS) simultaneously plotted po-sition at each sampling station, thus allowing geo-referencingof all measurements for each variable.

GPS tracks and dataflow information were used in a Geo-graphic Information System (GIS) to create highly detailedcontour maps of salinity and transmittance in relation tophysiographic features. Salinity was recorded as PSU andtransmittance on a relative scale of 0–5 units, with 5 beingideally transparent de-ionized water. The instrument systemwas used to locate point and non-point freshwater inputs andto determine spatial relationships of salinity and water clar-ity. Real-time data output discerned patterns in salinity dis-tribution and additional transects were then run in areas ofstrong gradients to enhance the spatial definition of theplume measurements. Discrete samples were taken from theflowing water stream during continuous sampling for labo-ratory analysis and instrument calibration.

Sediment Deposition

Five days after Hurricane Irene’s passage, we establisheda transect across Buttonwood Ridge just east of Taylor Creekto assess flood surge sediment deposition. Normally, the sur-face of this ridge is covered by a few centimeters of mangroveleaf litter (CORONADO-MOLINA 2000). After HurricaneIrene, a layer of carbonate sediment interspersed with sea-grass blades blanketed the ridge. Sediment depth was mea-sured from triplicate sediment cores collected at 5-m inter-vals along the transect. Flood surge depth was measured asthe level of the distinct mud line on the trunks of trees andshrubs across the ridge. By assuming a flat sea level during

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maximum stage, we were able to use the mud line measure-ments to map topography across the ridge.

Vegetation Sampling

Litterfall was collected from 10 litter traps on the Button-wood Ridge from 1996 through 2001. Each 0.25 m2 trap wasframed in wood and lined with mesh screen and were ran-domly placed throughout the ridge. Litter was collected fromeach trap on a monthly basis, sorted according to litter type,dried, and weighed. Averages and standard errors were cal-culated for each litter type and total litterfall at each site todetermine mass accumulation. We also estimated vegetativecover of the dominant seagrass species (Thalassia testudin-um) at multiple random sites in Little Madeira Bay. Coverestimates were made on a monthly basis using a quadrat-based, visual estimation of benthic area covered by seagrass.

RESULTS AND DISCUSSION

Creek Hydrodynamics

Storm events varied in both duration and intensity for thetime periods during which we evaluated the wetland-bay con-nection at Taylor Creek in the SE Everglades. Of the threeevents we studied, the ’96 winter storm was the longest induration (about 10 days), resulting in sustained, positive dis-charge from Taylor Creek into Little Madeira Bay during thefirst half of the event (highlighted region; Figure 2a). Evenwith minimal rains, salinity was low during this time as Tay-lor Slough water was pushed out into Little Madeira Bay bystrong northeast winds. When the winds subsided and shiftedto the southeast, flow reversed direction from positive to neg-ative discharge and stage increased nearly 0.3 m (Figure 2a).As bay water flowed back into Taylor Creek, salinity at themouth increased as much as 20 PSU, higher than it had beenthe previous five months. SUTULA et al. (2003) showed thatstrong northerly or southerly winds associated with thesetypes of events strongly influences discharge from TaylorCreek and other nearby mangrove creeks. Previous studiesin other micro-tidal systems around the Gulf of Mexico havedemonstrated that wind associated with frontal passages canaccount for significant fluxes of water and rapid changes instage (SMITH, 1977; PEREZ et al., 2000).

Unlike frontal passages, tropical storms and hurricanestypically bring substantial amounts of precipitation to southFlorida in addition to high winds. It is not uncommon for a1- to 2-day rain event associated with a tropical storm tobring 10 cm or more of precipitation to this region. This highprecipitation in the Everglades generally leads to highthrough-flow and subsequent changes in salinity regimesthroughout the estuarine ecotone.

T.S. Harvey passed directly over the SE Everglades, pro-ducing immediate and extended effects on the salinity andhydrodynamics of lower Taylor Creek. In Florida Bay, windand heavy precipitation associated with the passage of Har-vey led to an immediate increase in stage and negative dis-charge in Taylor Creek (Figure 2b). Overwhelmed by Ever-glades runoff, flow in Taylor Creek shifted direction and ex-hibited positive discharge as the storm passed (Figure 2b). In

1994, T.S Gordon produced a similar set-up (30 cm aboveaverage) in Florida Bay that did not return to pre-storm lev-els for 4 days (PITTS, 2001). Regional rainfall associated withT.S. Harvey (up to 30 cm) was great enough to sustain posi-tive discharge and produce a 15 PSU drop in salinity at themouth of Taylor Creek, lasting until the approach of Hurri-cane Irene 3 weeks later (Figure 2b).

After passing over the Florida Keys, the eye of HurricaneIrene also made landfall over the SE Everglades, showingstronger wind and precipitation effects. As with Harvey,southerly winds associated with Hurricane Irene produced aflood surge that reversed a pre-storm discharge of about 3 m3

sec –1 and forced bay water upstream into the Taylor Creeksystem at about 21 m3 sec21. Stage associated with the floodsurge peaked at 0.7 m above sea level as (Figure 2b). Afterthe center of the storm passed, strong northerly winds pre-vailed. This change in wind direction combined with fresh-water runoff resulted in high positive discharge (. 10m3.sec21) and a abrupt drop in stage (0.45 m over 8 hours).We observed the effect of this storm on discharge and salinitypatterns at the mouth of Taylor Creek for several weeks afterIrene passed across the peninsula and moved off into the At-lantic (Figure 2b).

A close sequence of storms such as these raises the ques-tion of how the synergy of combined events affects long-termecological dynamics in estuarine systems. The combinedevents we observed in 1999 had immediate and extended ef-fects on discharge and salinity patterns in Taylor River andLittle Madeira Bay (Figure 2). From 1 April 1999 up to thepassage of T.S. Harvey in late September 1999, daily salinityand discharge at the mouth of Taylor Creek averaged 19.4 69.7 PSU and 8.4 6 12.4 * 104 m3 d21, respectively. From thepassage of both Harvey and Irene through the end of 1999,daily salinity at this station was an order of magnitude lower,averaging 1.6 6 1.9 PSU, and mean daily discharge was morethan 3-fold greater (2.8 6 1.6 * 105 m3 d21) than the periodfrom April to September 1999. Furthermore, the total flux ofwater out of Taylor Creek during the last three months of1999 (2.6 * 107 m3) was more than 60% greater than the totalflux during the first 9 months of 1999 (1.5 * 107 m3), dueprimarily to the high precipitation associated with these twoclosely spaced tropical events.

Our findings suggest that the synergistic effects of theseclosely spaced storms on estuarine hydrology and waterchemistry was greater than if they had occurred more thanone month apart. Evidence of this phenomenon has been re-ported elsewhere during the same hurricane season. In 1999,Hurricane Floyd produced more than 20 cm of rain over anarea greater than 1000 km2 in the Chesapeake Bay water-shed. VALLE-LEVINSON et al. (2002) reported this eventresulted in at least 2000 m3 s21 of combined river dischargeinto the lower Chesapeake during the first day and a depth-independent outflow at the bay’s mouth. These authors pointout that Tropical Storm Dennis, which saturated the mid-Atlantic region less than two weeks prior to Hurricane Floyd,amplified the effect of Floyd in the Chesapeake watershed.

PAERL et al. (2001) noted a similar pattern of hydrologicalresponse to a three-storm sequence (Dennis, Floyd, andIrene) in Pamlico Sound (NC) also in 1999, concluding that

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Figure 3. Concentrations (mM) of Total (top) and Dissolved Inorganic (bottom) Nitrogen and Phosphorus at the mouth of Taylor Creek during the threeevents of interest (highlighted in gray). *No inorganic nutrient data were available from Tropical Storm Harvey.

these events had amplified and extended effects on phyto-plankton dynamics. During September and October 1999, theTar-Pamlico River Basin received approximately 96 cm ofprecipitation as a result of Hurricanes Floyd, Dennis, andIrene, resulting in an 8-fold increase in freshwater input toPamlico Sound. In the case of Dennis and Floyd, combinedriver discharge into the Chesapeake Bay was equivalent topeak spring discharge, but over a much shorter timeframe(VALLE-LEVINSON et al. 2002). These accounts clearly in-dicate the synergistic effects of these types of events on theannual patterns of estuarine function.

Creek Water Quality

Total nitrogen (TN) concentrations at the mouth of TaylorCreek exhibited no discernable trend before, during, or afterthe winter storm of 1996 (Figure 3). Concentrations of TNgenerally fluctuated between 50 and 70 mM during this pe-riod of time, which is a typical range for this time of year(CHILDERS, unpublished data). On the other hand, totalphosphorus (TP) concentrations increased more than 3-fold

from pre-event concentrations of 0.09 mM to approximately0.28 mM during the storm, and up to 0.42 mM immediatelyfollowing the event (Figure 3). The 10-day range in TP as-sociated with this November wind event was nearly equal tothe range in TP concentration measured during the entiremonth before and after the event.

Dissolved inorganic nitrogen (DIN; NOx2 1 NH4

1) and sol-uble reactive phosphorus (SRP) concentrations showed trendssimilar to TP, with increasing concentrations toward the endof the event and peak values shortly after the wind subsided(Figure 3). CAFFREY and DAY (1986) found increases in[NOx

2] in Fourleague Bay during frontal passages, reflectingthe high input from the Atchafalaya River, but we observedno such relationships for [NH4

1] or [SRP]. During the TaylorCreek events, [DIN] was less than 5% of [TN]. Similarly,[SRP] made up less than 5% of the total phosphorus fraction.These inorganic:total ratios for N and P are typical for TaylorCreek and therefore may not be event-related (DAVIS et al.,2001).

Prior to T.S. Harvey, [TN] near the mouth of Taylor Creek

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Figure 4. Total annual flux of total N and total P from Taylor Creekinto Florida Bay from 1996 to 1999. Hatched area in 1999 bars indicatethe flux of N and P that occurred only during the months of Septemberand October, the two months in which South Florida was affected byHurricane Floyd, Tropical Storm Harvey, and Hurricane Irene. * Includeonly fluxes measured from May 1 through December 31.

was approximately 65 mM, then declined to 45 mM, reachinga low of 40 mM a few days after the storm passed (Figure 3).[TP] increased from pre-storm levels of 0.28 mM to 1.16 mMafter the initial rains, then decreased by nearly an order ofmagnitude after the storm’s passage. We did not measureinorganic nutrients during Harvey; however, we collected dai-ly inorganic N and P data for 10 days immediately followingIrene’s passage. These post-hurricane data showed a 55% in-crease in [DIN] with peak discharge, paralleling an increasein [TN]. [SRP] increased noticeably (. 10-fold), fluctuatingbetween 0.06 mM and 0.13 mM for the remainder of Irene’swindow of effect (approx. 9 days). During this period, [SRP]made up as much as 40% of the [TP] in lower Taylor Creek,but [TP] remained low (0.15 to 0.3 mM) and constant follow-ing Irene’s passage (Figure 3). This high proportion of inor-ganic P may have resulted from the high precipitation andrunoff from this event. Flood surges originating from FloridaBay can also be an important source of carbonate sedimentand carbonate-bound phosphorus to this oligotrophic man-grove ecotone.

Until the passages of T.S. Harvey and H. Irene, rainfall insouth Florida was well below average for that time of year.This drought coincided with the La Nina phase of one of thestrongest ENSO events of the century. The contribution ofthese two events to the annual exchange of materials be-tween the mangrove zone and Florida Bay was explored us-ing total fluxes of TN and TP at the mouth of Taylor Creek.For the purpose of comparison, we estimated total annualfluxes for the years 1996 to 1999 (Figure 4). These datashowed fairly consistent exports of N and P from the TaylorCreek system to northeast Florida Bay across all years. How-ever, without Irene and Harvey, the export in 1999 wouldhave been considerably lower. We estimate that roughly 60–65% of the total N and P exported to northeast Florida Bayin 1999 was the result of these two storm events (Figure 4).A similar approach taken by PAERL et al. (2001) indicatedthat the sequence of storms affecting Pamlico Sound in 1999accounted for a substantial 2-month (September and October)load of DIN into the estuary, which was approximately 70%of the annual average DIN load from 1994–1997. Findingssuch as these underscore the importance of these types ofpulsed events in the exchange of materials at the coastalmargin.

Bay Water Quality

High resolution mapping of water quality showed that sa-linity in Little Madeira Bay was considerably lower followingthe passage of Irene than in the same month of the followingyear—a wet season period with no tropical storm activity(Figure 5a). The influence of Hurricane Irene on the physico-chemistry of the Everglades-Florida Bay transition at LittleMadeira Bay was unambiguous. Immediately following Irene,salinity ranged from 0 to 3 PSU in Taylor Creek and 1 to 9PSU throughout Little Madeira Bay. The magnitude ofIrene’s effect is obvious with low salinity water extending faroutside of Little Madeira Bay into Florida Bay (Figure 5a).On 19 October 2000, salinity ranged from 10 to 21 PSU inLittle Madeira Bay, with low salinity water confined to the

mouth of Taylor Creek (Figure 5a). In both periods, salinityin Taylor Creek was , 5 PSU, which is typical for the wetseason. No high resolution spatial data are available for theperiod immediately prior to Hurricane Irene, but Little Ma-deira Bay salinity in the beginning of the Atlantic hurricaneseason (June 1999) ranged from 23 to 33 PSU, thus demon-strating ‘‘pre-event’’ conditions were considerably more sa-line.

In both October 1999 and October 2000, the transparencyof Taylor Creek was high (80 to 90% relative to distilled wa-ter; Figure 5b). Furthermore, in almost all cases, the waterin Taylor Creek was clear and tannic while Little MadeiraBay and adjacent Florida Bay were generally more turbid.Following Irene, high discharge produced plumes of clear, lowsalinity slough water that extended from the mouth of TaylorCreek into Little Madeira. This plume effect was clearly ev-ident in the 1999, where lowest salinities coincided withclearest waters of the northern half of the bay and highestsalinities (e.g. in the far south east corner of Little MadeiraBay) coincided with more turbid bay water (Figure 5b).

During a storm event such as Irene, with high winds andrainfall, opposing forces affect water clarity. High winds leadto re-suspension of bay sediments, while the introduction ofclear freshwater from the Everglades reduces turbidity. Re-sults of this process can be seen in the spatial pattern ofwater clarity in the days following Irene, with the southernhalf of Little Madeira Bay and portions of Florida Bay char-acterized by lower transparency and Taylor Creek and thenorthern half of Little Madeira Bay showing high transpar-ency. In the quiescent period of October 2000, the entire areawas typified by water of high transparency, likely a result oflow wind-wave energy and wet season freshwater inputs dur-ing this time (Figure 5).

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Figure 5. Contour plots of surface water salinity (left panels; A) and transparency (right panels; B) in Little Madeira Bay and nearby Florida Bay duringOct 1999, soon after Hurricane Irene (top panels; 1), and in October 2000, during a period without storms (bottom panels; 2).

Figure 6. top: Deposition of carbonate sediment (mean 6 std dev) alonga transect across the Buttonwood Ridge (near Taylor Creek) from Hur-ricane Irene (10/15/99). bottom: Water depth profile along transect (esti-mated from mud line on vegetation), indicating surface topography ofButtonwood Ridge. Dashed line represents surface water level duringmeasurement.

Sediment Deposition

The Buttonwood Ridge is a depositional berm separatingdwarf mangroves and sawgrass marshes of the SE Ever-glades from direct marine influence of Florida Bay and theGulf of Mexico (see image in Figure 1). Only during strongwind/precipitation events such as Hurricane Irene do waterlevels exceed ridge height, leading to flow over this embank-ment. During Irene’s passage, the flood surge was highenough to inundate the Buttonwood Ridge, depositing baysediments in the mangrove forest. This flood surge ranged

from about 20 to 50 cm in depth over the ridge—based onheight measurements of the visible, fresh mud line on thetrees along our transect—and deposited as much as 5 cm ofcarbonate sediment on the ridge (Figure 6). This depositionwas confined to a 60-meter zone in the center of the ridge. Itappears that flow rates were too strong for deposition to occurat the leading edge of the ridge. We suspect that sedimentdeposition occurred once the initial surge water velocities de-creased and the floodwaters, with their bay-derived sedimentload, receded back across the ridge after the storm’s passage.

Sediment deposition was variable across the ButtonwoodRidge, possibly as a function of the ridge’s micro-topographyand the accumulation of sediments in depressions as thesurge receded (Figure 6). Preliminary data from Surface El-evation Tables (SETs; BOUMANS and DAY, 1993) in the Ev-erglades estuarine ecotone indicate inter-annual soil eleva-tion changes of only 5–15 mm (SKLAR, unpubl. data). Hur-ricane Irene deposited several times that amount on the But-tonwood Ridge—in one single event. The implications of thistype of storm-related deposition have received little treat-ment. However, this may be an important process allowingthis system to accrete vertically, answering rising sea level.Further, the structure of the Everglades mangrove ecosystemmay be dictated by these kinds of storm events, which deliversediments, nutrients (especially phosphorus), and water tothe region in pulses.

Vegetation

In shallow coastal areas, reductions in water temperatureand light availability associated with winter storms havebeen shown to exert strong controls over seagrass survival(CABELLO-PASINI et al. 2002). Seagrass cover data werenot available for 1996 or 1997 to substantiate this. The pro-duction of mangrove litter showed no noticeable effect of the1996 windstorm or of Tropical Storm Harvey (Figure 7). How-ever, Hurricane Irene produced a noticeable pulse of man-

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Figure 7. Monthly values for mean leaf litterfall (6std error) for man-groves along the Buttonwood Ridge of Taylor Creek. Timing of ’96 winterstorm and Hurricane Irene passage indicated.

Figure 8. Mean density of Thalassia testudinum short shoots in LittleMadeira Bay from June 1998 to August 2000. Timing of Hurricane Irenepassage indicated.

grove leaf litter (. 6 g dry weight m22) for the month of Oc-tober 1999 relative to other months for which we have pro-ductivity data. The flood surge and subsequent sediment de-livery associated with Irene’s approach also led to thescouring of Little Madeira Bay and coincident reduction inseagrass cover. We observed a decline in seagrass cover from.250 short shoots m22 to 0 m22 at bay sites near the mouthof Taylor Creek before and after Hurricane Irene’s passage(Figure 8). Further evidence of this was found in the newsediment deposited on the Buttonwood Ridge, which includednumerous green seagrass blades.

The pulse of mangrove and seagrass detritus generated byhurricane Irene may have contributed to the greatly elevatedcreek water concentrations of TN, SRP and DIN we describedearlier (Figure 3). Rapid leaching of newly deposited littercan account for a significant flux of labile organic matter andnutrients to the water column (BENNER et al., 1986; DAVISet al., 2003), and may represent a significant source of nutri-ents to the mangrove ecotone following storms of this mag-nitude. Evidence from a study in coastal Louisiana supportsthis, as a hurricane-induced pulse of litter led to significantchanges in water quality (RYBCZYK et al., 1995).

SUMMARY

Coastal ecosystems are influenced by pulsed events thatoperate across a range of spatial and temporal scales. Global-scale events such as El Nino operate over long timescales,affecting annual hydrologic and productivity patterns (ZIM-MERMAN and ROBERTSON, 1985; CHILDERS et al., 1990).Short-term events such as hurricanes may last from hours todays, but can have both acute and lasting effects on ecosys-tem processes (SMITH et al., 1994; PAERL et al. 2001). Anestuary’s response to a rapid series of these types of stormevents can reveal a great deal of information about its resil-ience as well as the importance of disturbance in shapingecosystem structure and function (PAERL et al. 2001). Suchresponses can be observed through continuous monitoring

programs (remote or in situ) or through intensive ecologicalsampling that overlaps a storm event’s window of effect. Thelatter are often lacking due to the logistical problems in as-sembling a study immediately following an event or the dif-ficulty in initiating a sampling study that coincides with thetiming and location of an event’s strike.

We were fortunate in that our continuous monitoring andlong-term sampling programs in Taylor Creek coincided witha number of frontal winter storms, tropical storms, and hur-ricanes. During our study period (1995–2001), the SE Ever-glades mangrove ecosystem was affected by a number of trop-ical and winter storm events. These events resulted in dra-matic changes in surface water movement and chemistry inTaylor Creek and Little Madeira Bay, sediment delivery tothe Buttonwood Ridge, increased litterfall, and scouring ofseagrass beds. In many years, these kinds of events seem toaccount for a substantial proportion of the annual flux offreshwater and materials between the SE Everglades andFlorida Bay. Depending on a storm’s path, events such asHurricane Irene may serve as periodic sources of carbonatesediment and carbonate-bound phosphorus to this P-limitedecotone, thus contributing to the maintenance of the man-grove wetland’s productivity and position relative to sea level.

ACKNOWLEDGEMENTS

The authors wish to thank the folks who graciously helpedout in the field and with data analysis, including Martha Su-tula, Brian Perez, Steve Kelly, Chelsea Donovan, and Mi-chael Korvela. South Florida Water Management Districtand the National Park Service, especially employees of theEverglades National Park Key Largo Ranger Station, wereinstrumental in making this work possible. This research wasfunded by SFWMD contracts #C-E6608 to JWD and #FR-976to DLC. This material is also based upon ongoing work in theFlorida Coastal Everglades Long-Term Ecological Research(FCE-LTER) Program and supported by the National ScienceFoundation (DEB9910514).

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