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Gymnodinium breve red tide blooms: Initiation, transport, and consequences of surface circulation



From its source waters in the Gulf of Mexico the led tide dinoflagellate, Gymnodinium breve is moved throughout its oceanic range by major currents and eddy systems. The continental shelf off the west coast of Florida experiences frequent G. breve blooms (in 21 of the last 22 years) where the spatially explicit phases of G. breve blooms are closely coupled to physical processes. Bloom initiation occurs offshore and in association with shoreward movements of the Loop Current or spinoff eddies. A midshelf front maintained by seasonal wind reversals along the Florida west coast may serve as a growth and accumulation region for G. breve blooms and contribute to the reinoculation of nearshore waters. Local eddy circulation in the northeastern Gulf of Mexico and in the Dry Tortugas affects the retention and coastal distribution of blooms while the Florida Current and Gulf Stream transport cells out of the Gulf of Mexico and into the U.S. South Atlantic Eight. The causes of bloom dissipation are not well known but mixing or disruption of the water mass supporting G. bp-eve cells, especially in combination with declining water temperatures, are important factors.
Gymnodinium breve red tide blooms: Initiation, transport, and consequences
circulation of surface
Patricia A. Tester
National Marine Fisheries Service, NOAA, Southeast Fisheries Science Center, Beaufort Laboratory, Beaufort, North
Carolina 28516-9722
Karen A. Steidinger
Florida Department of Environmental Protection, Florida Marine Research Institute, 100 Eighth Ave., SE, St. Petersburg,
Florida 33712
its source
in the Gulf of Mexico the red tide dinoflagellate,
Gymnodinium breve
is moved throughout
its oceanic range by major currents and eddy systems. The continental shelf off the west coast of Florida experiences
frequent G. breve blooms (in 21 of the last 22 years) where the spatially explicit phases of G.
blooms are
closely coupled to physical processes. Bloom initiation occurs offshore and in association with shoreward move-
mcnts of the Loop Current or spinoff cddics. A midshelf front maintained by seasonal wind reversals along the
Florida west coast may serve as a growth and accumulation region for G.
blooms and contribute to the
reinoculation of nearshore waters. Local eddy circulation in the northeastern Gulf of Mexico and in the Dry Tortugas
affects the retention and coastal distribution of blooms while the Florida
and Gulf Stream transport cells
out of the Gulf of Mexico and
the US. South Atlantic Bight. The causes of bloom dissipation are not well
known but mixing or disruption of the water mass supporting G. breve cells, especially in combination with declining
water temperatures, are important factors.
Much of what is known about the distribution of the toxic
dinoflagellate Gymnodinium breve Davis [ = Ptychodiscus
brevis (Davis) Steidinger] is explained by oceanic circulation
patterns. G. breve cells are positively phototactic (or nega-
tively geotactic) (Steidinger 1975; Heil 1986) and can con-
centrate in the upper water column during the day. There
they behave like surface drifters, only smaller writ. The res-
ident population is in the Gulf of Mexico and G. breve is
transported throughout its range by the Gulf Loop Current,
the Florida Current, and the Gulf Stream, with the warmth
of the Gulf Stream fostering this subtropical species as it
moves north of 31”N. G. breve has been recorded throughout
the U.S. South Atlantic Bight (Tester et al. 1993) and be-
G. breve rarely occurs in shelf waters north of Cape Hat-
teras, North Carolina (Marshall 1982; see also Churchill and
Cornillon 1991), but the Gulf Stream may carry it farther
(see Fraga and Sanchez 1985). A drift bottle released in an
October 1966 study off the central west Florida shelf was
recovered in the Outer Hebrides a little over a year later;
another released in February 1967 reached Belgium in 187
d (averaging >50 km d-l) (Williams et al. 1977). So it seems
likely that long before
1497, when Vespucci, credited
We thank S. Baig, who supplied the Gulf Stream Frontal Anal-
yses (GOES) and is a continuing source of information and support.
T. Lee is acknowledged for his ideas on the entrainment of G.
cells in the Tortugas Gyrc. J. Turner, G. Cervetto, M. Geesey, C.
Lewis, and A. Smith helped draft figures. E. Haugen and three re-
viewers made valuable comments on the manuscript. B. Roberts
maintains and shared the G. breve bloom log. R Stegmann and K.
Carder provided CZCS imagery. The source of the AVHRR image-
is B. Stone, NESDIS and the NOAA Coastwatch Program.
with being the first European to explore the Gulf of Mexico,
sailed out of the gulf in the direction of the “maestrale” for
870 leagues to Cape Hatteras and then turned eastward to-
ward Bermuda before returning to Spain (Galtsoff 1954), G.
breve had made a similar voyage.
The reports of discolored water and effects of phycotoxins
in tropical Atlantic waters were recognized and recorded in
ships’ logs by 1530-1550 (Martyr 1912). As early as 1844
popular accounts of G. breve blooms were linked with nox-
ious “gases” and massive fish kills along the west coast of
Florida (Feinstein et al. 1955). Prophetically, in that same
year an intensive study OF ocean currents was begun by Mau-
ry (1859). Data on ocean circulation, physical processes, and
the distribution and biology of G. breve would accumulate
for more than a century before this information would be
coupled, the consequences appreciated, and a model concep-
tualized to help focus research efforts,
Although Lackey (1956) reported G. breve from Trinidad
in the southern Caribbean basin (cell counts were not veri-
fied), there were no recorded cases of neurotoxic shellfish
poisoning reported (S. Hall pers. comm.), and there have
been no subsequent observations of G. breve anywhere in
the Caribbean. Early interest in the circulation of the Carib-
bean, though, stemmed from its role as the source region for
water flowing into the Gulf of Mexico. In this semi-enclosed
basin the eastern gulf is characterized by anticyclonic cir-
culation and is dominated by two currents. The Yucatan Cur-
rent (75 cm s-l), entering between the Yucatan Peninsula
and Cuba, becomes the Loop Current as it extends northward
into the gulf and returns southward along the west Florida
1040 Tester and Steidinger
Fig. I. Generalized surface circulation of the Gulf of Mexico. Arrows denote steadiness of
current drift. (Redrawn from UNEP/CEPAL Caribbean Environment Program, Project 1037.)
continental shelf. It exits the gulf between the Dry Tortugas
and Cuba where it is known as the Florida Current (165 cm
s-l) (see Hofmann and Worley 1986) (Fig. 1). The extent of
northward penetration of the Loop Current (Maul 1977;
Vukovich et al. 1979), its spinoff eddies (Dietrich and Lin
1994; Sturges 1994), and its intrusions onto the west Florida
continental shelf vary seasonally (Huh et al. 198 1) and great-
ly affect the potential of bloom initiation, transport, and re-
tention (Haddad and Carder 1979; Haddad 1982; Lee et al.
1994). Rotating eddies can be shed from the Loop Current
and propagate westward across the gulf (Liepper et al. 1972;
Maul and Vukovich 1993). Cross-basin surface transport
also has been documented in drift-bottle studies (Williams
et al. 1977). Summer-fall bottle releases from the west Flor-
ida shelf frequently were recovered from Texas beaches
(Matagorda to Brownsville) to Vera Cruz, Mexico. The gen-
eral circulation patterns of the south central and western Gulf
of Mexico are clockwise but the velocities are less intense
(15-25 cm s I). The only exceptions to this general clock-
wise pattern are the cyclonic flows in the extreme north-
eastern and northwestern areas of the gulf (Molinari 1980;
Vastano et al. 1995)-regions where elevated background
concentrations of G. breve cells (>lOO cells liter ‘) have
been noted (Geesey and Tester 1993).
Throughout the Gulf of Mexico and the U.S. South At-
lantic Bight, G. breve is found in background concentrations
(l-1,000 cells liter- ‘) except in areas off the Texas coast and
the west Florida coast where local circulation may play a
role (Fig. 2; see Geesey and Tester 1993; Tester et al. 1993).
Although G. breve blooms have occurred in many different
areas in the Gulf of Mexico, from Yucatan in the south (Gra-
ham 1954), to the lower Laguna Madre, in the Mexican state
of Tamaulipas in the western gulf (Gunter 1952; Wilson and
Ray 1956) to Freeport, Texas (Burr 1945; Trebatoski 1988),
and around the northern gulf coast, they are most frequent
along the west coast of Florida. Blooms there are especially
frequent from Clearwater to Sanibel Island (Joyce and Rob-
erts 1975; K. A. Steidinger and B. S. Roberts unpubl.), oc-
curring in 21 01: the last 22 years. These blooms on the
southwest Florida shelf serve as a source for cells inoculat-
ing the U.S. South Atlantic Bight (Murphy et al. 1975; Stei-
dinger et al. 1995; Tester et al. 1991).
Bloom initiation
The regions of the GulF of Mexico that experience blooms
of G. breve lasting more than 2 months include the west
Florida shelf (C.earwater to Sanibel Island), the Campeche
Bay between Rio Ciatzacoalcos and Rio Grijalva (Smithson.
Inst. 1971), and the Texas coast between Port Arthur and
Galveston Bay. 411 have common features conducive to the
formation of blooms. Each of these areas is adjacent to a
continental shelf break where it intersects with the perma-
nent seasonal thl:rmocline (Fig. 3). The isothermal water on
the outer shelf results in a minimum bottom temperature of
20°C in areas off Texas and west Florida (NOAA 1985) and
Campeche Banks. Consequently these areas may provide an
important winter refuge for G. breve. These same areas also
experience either persistent, intermittent, or event-related
slope-shelf upwelling. This is best known for the west Flor-
ida shelf where blooms can occur any time of the year but
are typical in late summer and fall when >70% of the out-
breaks have begun. Bloom concentrations first appear OFF-
shore (Dragovich and Kelly 1966; Steidinger 1975; Steidin-
ger and Haddad 1981) and are associated with the fronts
caused by the onshore-offshore meanders of the Loop Cur-
rent water along the outer southwest Florida shelt Water on
Fig. 2. Gymnodinium breve nonbloom, background concentrations in cells liter-‘. Samples were
taken during 1989-1991 by ships of opportunity (after Gcesey and Tester 1993; Tester et al. 1993).
Fig. 3. Winter bottom-water temperatures on the continental shelf of the Gulf of Mexico basin
(redrawn from NOAA 1985). Regions of persistent upwelling off the southwest Florida shelf, Texas-
Louisiana coast, and the Yucatan Peninsula are indicated by the arrows.
Tester and Steidingrr
Fig. 4. CZCS image. 14 November 1978, providing an estimate of chlorophyll a in &.g liter
(inset) during a Gymnodinium
bloom off the west Florida coast from soulh of Tampa Bay to
the Florida Keys. When processed ar a higher
chlorophyll a is detectable at G.
cell concenlraliuns that are not viqihlc to the human eye (i.e. Stump Pass, top arrow. 6.7X IO’ cells
hoer ‘; Doctor-Gordon Passes, hottom anow, 4.6X10‘ cells
the leading edge of an eddy or meander is generally sinking
and that on the trailing edge is rising, until the feature ex-
periences bottom drag (Dietrich and Lin 1994) and forms a
midshelf front.
Fronts represent a dynamic area of nutrient regimes and
light conditions which can favor accumulation and growth
(<l div. d ‘, often 0.2-0.5 div. d-‘) of dinoflagellates.
Bloom species such as G.
cf. aura-
lum (European waters) are well adapted to such environ-
ments and can grow throughout the euphotic zone. They
have a high photosynthetic capacity at low light and are
light adapted at varying intensities (Shanley 1985; Garcia
and Purdie 1992) although photoinhibition thresholds are
species-specific. Once growth occurs, it takes 2-X weeks to
develop into a hloom of fish-killing proportions (1-2.5X IOx
cells liter-‘) depending on physical, chemical, and biological
Some species such as G.
breve, G.
cf. aureolum, and
gulodinium plydrum
have growth and competitive exclu-
sion strategies that can lead to almost monospecific surface
blooms of these bpecieb (as biomass) (Sttidinga and Vargo
Morin et al. 1989). Such blooms (G.
breve, G.
aureolum) can cclvei- a surface area of up to 1.4-3.0x10’
km’ (Steidingcr and Joyce 1973; Holligan 1985; Vargo et al
1987) and although biomass concentration is patchy, chlo~
rophyll a values from >2 to > 100 mg tr’ make the rest&
tant discolored surface water detectable by color sensors 011~
board satellites (Fig. 4). In the case of G.
the CZCS
sensor detected chlorophyll a from cells at densities one-
two orders of magnitude less (IO”-1 Oi) than are present when
discolored water is detectable by the human eye (>lO”) (K.
Haddad and K. Steidinger per% comm.).
Both G.
and G. cf. auwulum discolor surface waters
and are phototactic; in daylight hours cells are at or near the
surface; at night they are dispersed (Hollignn 1985; Heil
1986; Geesey and Tester 1993). In addition to their ability
to exploit light regimes, both species have advantages in
nutrient dynamics. Both assimilate nitrogen at low light and
are able to utilize organic as well as inorganic nutrients (Var-
go and Shanley 1985; Steidinger and Vargo 1988; Dahl and
Tangen 1993; Shimizu et al 1995). When G.
have been tracked, the zone of initiation (cell numbers
>I,000 liter ‘) develops from 18 to 74 km offshore (St&
dinger and Haddad 1981) and the strongest evidence for this
lies in the cell distribution along cross-shelf transects sam-
G. breve blooms 1043
Cells ‘I Id liter”
a > 400
igj 200 - 400
[7 >5-200
0 -=5
Fig. 5. Spatial and temporal distribution of Gymnodinium breve cell abundance along the west
Florida coast during a late summer-fall 1976 bloom.
pled by personnel from the Florida Marine Research Institute
(FMRI) and Mote Marine Laboratory.
The first example is taken from transects sampled during
a late summer bloom in 1976. A series of stations from the
inlets or nearshore to 32-70 km offshore was made between
Distance from Shore (km)
Fig. 6. Distribution and abundance of Gymnodinium hew cells
along cross-shelf transects north of Boca Grande Pass to >66 km
offshore taken bctwcen 24 September and 11 October 1976.
22 September and 15 October. Concentrations above back-
ground were first noted offshore between Sarasota and Boca
Grande Pass in late September (Fig. 5). During the following
week in the same area, cell concentrations increased and then
generally spread south. No nearshore or inlet samples during
this time were either positive for G. breve or above back-
ground until 2 weeks later. Data from north of Boca Grande
Pass are representative of the cell distributions from transects
run perpendicular to shore (Fig. 6). Typically the cells are
moved from the midshelf to onshore and then, under the
influence of the wind and(or) alongshore currents, move up
or down the coast (Fig 5). From 7 to 11 October there were
still high numbers (>l-6X 10s cells liter -I) of G. breve 3-
25 km offshore north of Boca Grande Pass. The bloom had
moved south and was parallel to the coast with lo-IOO-fold
lower cell concentrations onshore than offshore. By 13-14
October the bloom was centered south of Boca Grande Pass
and had intensified. Counts offshore of Sanibel Island were
10” cells liter-‘; counts nearshore ranged frown 1.5 to
70X lo1 cells liter-‘. Farther south at Naples there were no
cells in the 25&m station, but nearer shore cells were noted.
After 15 October the inshore passes between Clearwater and
Naples had < 1,000 cells liter -I and sampling was suspend-
A similar pattern of bloom development and movement is
evident from blooms in November-December 1979 and Sep-
I 044
Tester and Steidinger
Cells I 10’ liter“
5 350
fl 200 - 350
l <5
Fig. 7. Spatial and temporal distribution
of’ Gymnodinium breve
cell abundance along the weat
Florida coast during late fall 1979.
tember-November 1985 (FMRI data). The 1979 bloom was
first reported 130 km northwest of Clearwater and the ear-
liest cell counts from the Clearwater and Cedar Key areas
(26-30 November) indicated that it was a large bloom (Fig.
7). Note however, there were no cells or very low counts
(O-2,000 cells liter-‘) onshore until I O-l I December, when
offshore numbers were dropping and the bloom seemed to
have moved south loo-120 km and onshore. The third ex-
ample is from a 1985 bloom which was first sampled 16 km
off Cedar Key and southward (101-10s cells liter -I> on 10
September (Fig. 8). Again the passes and nearshore were
free of G.
cells until more than a month later. The
alongshore (south) and onshore movement of the bloom is
evident; the offshore stations between Clearwater and Sar-
asota were cell-free during the first sampling period, but l-
2 weeks later there were up to lo”-10“ cells liter-‘. Within
4-5 weeks between lo5 and 10” liter-’ were observed on-
shore and south of Sarasota and Boca Grande Pass (4 No-
Bloom transport
There is evidence that some blooms can be maintained
within the midshelf zone and continually inoculate the near-
shore waters or recur in a “high occurrence zone” from
Clearwater to Sanibel Island (Steidinger and Roberts un-
publ.). One possible mechanism for this is the circulation
pattern reported by Weisberg et al. (1996). They describe
seasonal wind reversal, (northeast-southwest flow) on the
midshelf that result in zero mean Rows both in the along-
shore and crossshore directions during a 16-month period.
However, the mcnthly means can be re.latively large, and
Weisberg et al. suggested the maximum values have a bar-
oclinic origin via a thermal wind relationship. Recent re-
search on the west Florida shelf circulation describes two
basic patterns, a summer pattern (April-September) and a
winter pattern (October-March) characterized by a semiper-
manent anticyclojiic eddy (Weisberg et al. 1996) on the
northwestern Florida shelf in the Apalachee Bay-Middle
Grounds area (H. Yang and R. Weisberg :pers. comm.) This
feature dominates the northeastern shelf onshore of the 50-m
isobath (Yang and Weisberg pers. comm) and may be re-
sponsible for the ~entrainment and transport of cells north-
ward to the Florida panhandle. Haddad ( 1982) recorded an
example of a red tide preceded by the shoreward advection
of a bottom thermocline between 10 August and 3 Septem-
ber 1978 when the bloom surfaced at 19 km from shore.
Similar transport along a thermal gradienit may explain the
spring-summer bloom of 1995 when a G.
red tide
apparently moved north up to Cedar Key and then onto the
Florida panhandle 55-l. 10 km offshore and subsequently in-
oculated inshore waters (FMRI unpubl. data).
From Tampa Bay south small-scale eddy features (<lo0
km; Maul 1977; Hela et al. 1955) or filaments (R. Stumpf
pers. comm.) may also play a role in the translocation of
offshore blooms. Frontal eddies (Loop Current water) and
G. breve
ca Grande Pass
anibel Island
Fig. 8. Spatial and temporal distribution of
Gymnodinium breve
cell abundance along the west
Florida coast during a late summer-fall bloom in 1985.
on-offshore meanders of the Loop Current move southward
along the outer southwest Florida shelf (every 2-14 d) (Pal-
uszkiewicz et al. 1983), and there is evidence for a south-
ward mean flow over the shelf when the Loop Current is at
the shelf edge (Sturges and Evans 1983). The annual cycle
of wind stress, northward during summer and southward in
fall, is responsible for the persistent upwelling (summer) or
downwelling (fall) found over the west Florida shelf (Lee
and Williams 1988) and may concentrate or disperse blooms
depending on the site and timing of the bloom.
Another recently described eddy system operative be-
tween the Tortugas and the upper keys is dependent on a
well-developed Loop Current and the consequent offshore
position of the Florida Current. The Tortugas Gyre is a cy-
clonic recirculating feature (100-l 80 km) with a duration of
40-J 08 d that has a strong influence on the transport and
retention of zooplankton and larval fish in the lower Florida
Keys (Lee et al. 1994). In the intervals between gyre recir-
culation periods there are episodes (20-30 d) of intense east-
ward flow (Lee et al. 1994). This feature may provide insight
into the distribution and transport of G.
From the west
coast of Florida to the Atlantic in 1994-1995 (T. Lee pers.
comm.). The bloom started in September 1994, between
Tampa Bay and Sanibel Island off the west coast of Florida.
In January and February 1995, extensive fish kills were re-
ported in >5,000 km2 of open water westward from the Flor-
ida mainland to the Dry Tortugas; a sample with 1.5X lo5
cells liter-’ was counted from Sanibel on 1X January. In
February, coincident with fish kills off the southwest Florida
coast, a sample from 12 km off Duck Key (14 February
1995) contained 9.6X 10” cells liter-’ (Fig. 9). Concern about
the possibility of G.
transport to the Atlantic prompted
sampling off West Palm Beach from September 1994
through early March 1995. Cell counts at West Palm Beach
were only O-6 cells liter-’ from September to December
1994 but from 17 to 24 February 1995, as the Gulf Stream
impinged on the West Pplm Beach sampling site and formed
a meander, G.
cell densities increased to -2X lo4 cells
liter-’ (Fig. 10) (Steidinger et al. 1995). As the Gulf Stream
moved offshore, away From the sampling area, cell numbers
decreased (27 February-3 March). From 3 to 7 March 1995,
samples from the Occulina Reef National Park (-27”53’N,
79”58’W) off Cape Canaveral, north of the West Palm Beach
site contained O-80 cells liter-.‘, but the Gulf Stream did not
impinge on the reef during the 4-d sampling period. Fourteen
days after the Gulf Stream meander migrated past West Palm
Beach, G.
cells were found in 10 of 12 samples from
the outer shelf of Onslow Bay, North Carolina (33”51’N,
76”53’W, 23.4’), >l,OOO km downstream. Although the cell
counts were low (55 cells liter .I) the presence of G.
in >80% of the samples is unusual in early March. Fortu-
nately, the dire oceanic menace (as depicted by the
on 22 February 1995) following the course of the
Gulf Stream was exaggerated. In early March the inner shelf
water of Onslow Bay is too cold (<12”C) to support the
growth of G.
and wind mixing prevented water-col-
umn stabilization conducive to bloom development.
Prior to 1969, G.
had not been recorded in the U.S.
Tester and Srcidiqer
South Atlanuc Bighl, but Lackey (I 969) found one cell in
an unpreserved sample taken from Boa Katon during a wa-
ter-quality study. Nol until autumns 1972, 1976, and 1Y77
after the typical manifestations of G. hrrve blooms (e.g. eye
and respiratory irritation, fish mortality) were described by
beachgoers from Miami to Palm Beach (Murphy et al. 1915;
Robens 1979), did we undcrsland that blooms--even shorl-
lived ones (t I month)-could occur outside the Gulf of
Mexico basin. These first blooms recorded in the U.S. South
Allantic Bight were restricted in area and intcnsily. No cells
were found a\ far north as Cape Canaveral. and the count’:
wcrc neither hrgh nor persistent (2-100X 10’ cells liter ‘).
The eveas were conxdered the result of concentration and
transport of cells to the east coast by unusual Loop Current
parterns (Murphy et al. 1975).
The seasonality of the west Florida shelf circulation and
wind liclds alho contributes to the likelihood of shelf water
being advected into the Florida Current for transport to the
U.S. South Atlantic Bight (Williams et al. 1977). Late sum-
mcr-aulumn blooms have the greatest potential for transport
to the Atlantic coast bccausc summer transport rates are the
highal and “detrainment” is greatest in Pall due to low,
inconsistent trampor, (Maul and Bravo I9XY). The thrw jr-
cas of dcuainment identified by Maul nnd Bravo (19X9) ah
likely was for receiving flotsam and .jetsnm are southeast
of Cape Canaveral, eat of St. Augu\tinc, and southeast of
Onslow Bay. G. brew was TV prove itself as an apt surface-
drifter and a good test of their ideas.
Cells from a May-October 1980 war Florida shelf bloom
were transpor~cd farther than any recorded up to thai lime.
Gulf Stream Frontal Analyses (compobite GOES satellite im-
agwy) confirmed a warn-water intrusion 7-10 November
off Jacksonville (Fig. I I) and by 14 Novcmbcr local Jack-
sonville residents were suffering sore throats and watery
eyes; by 2528 November G. brew counts were 6.7X IO‘
cells lite: and the bloom had spread >I00 km south LO
Daytona where beachgoers were affected by exposure to the
surf (FMKI data). Cells were found off Cape Canaveral
(Melbourne) on 5 Decemhcr and between Jacksonville and
Cape Canaveral mcandcrs of warm water shoreward of the
western edge of the Gulf Stream were evident in GOES in-
agery of 8 December 1980 (not shown).
The 1980 east coast G.
red tide dcmonaated that
Gulf Stream meanders off Jacksonville could inoculate in-
G. breve blooms 1047
14 15 16 17 23 24 25 2 3 6
Feb 95 March 95
10. Gymnodinium breve
cell counts near West Palm Beach, FL in relation to the position of the Gulf Stream. When the western
front of the Gulf Stream was closer to the ncarshore sampling site the cell counts were I-18.8X lo3 cells liter .I. When the Gulf Stream
was seaward of the sampling site G.
cell concentrations were <IO0 cells
~I (after Steidinger et al. 1995). Cape Canaveral is the
cape immediately north of West Palm Beach (0).
shore areas
and be transported by countercurrents south
along the coast beyond Cape Canaveral. In 1983, another
transport occurred concurrently as a meander of the Gulf
Stream Formed south of Jacksonville on 10 October (S. Baig
pers. comm.). In the area from Daytona to below Cape Ca-
naveral (Volusia and Brevard Counties) all three signs of a
G. breve red tide were evident (i.e. human respiratory irri-
tation, fish kills, and discolored water). The cell counts were
much higher than for previous east Florida shelf blooms
(5.5X106 cells liter I, 10 October). The following day cell
counts were 1X 10’ cells liter ‘; during the next 10 d, red
tide affected areas immediately south of Cape Canaveral
(Patrick Air Force Base, Cocoa Beach, and Melbourne). By
28 October, it had moved 55 km south of Cape Canaveral
(Sebastian Inlet) and remained there until 4 November. By
23 November, the red tide had dissipated and shellfish har-
vesting areas were opened.
Prior to that east coast event, a well-developed red tide
was detected on the west coast on 6 October. Patches of dead
fish and surface-water discoloration (indicative of cell counts
23 X 10” cells liter ‘) were reported from Sarasota to Venice
from shore to 15 km offshore. Inshore G. breve concentra-
tions were 4.5 X 10” cells liter-’ and beachgoers were expe-
riencing respiratory irritation. Cell counts from 7 October
were 3X 10” cells liter I 9 km off Captiva Island in the Char-
lotte Harbor area. This is the classic source area for transport
of red tides from the west coast to the east coast of Florida.
Because it takes 2-8 weeks for a red tide to develop con-
centrations of +2.5X lo5 cells liter offshore, this west Flor-
ida bloom was the likely source for cells inoculating the
Jacksonville area.
Gulf Stream transport also was implicated in an unusual
G. breve fall and winter (19X7-1988) bloom along the coast
of North Carolina which continued for 4-5 months (Tester
et al. 1991). Thirty days before this bloom there was a late
summer bloom off the southwest coast of Florida. The cou-
pling of these two events is supported by transport time of
an Argos-tracked surface drifter (60 km d I) making the
same passage in -20 d (Ortner et al. 1995) and the drift
bottles recovered from Wrightsville Beach, North Carolina,
between 3 1 and 100 d after their release off the west Florida
shelf (Williams et al. 1977). The continental shelf between
Cape Hatteras and Cape Lookout where this bloom occurred
is the narrowest of any in the U.S. South Atlantic Bight north
of the Miami area and is frequently overwashed by meanders
of Gulf Stream water, some of which nearly reach the barrier
1048 Tester and Steidinger
Fig. Il. Shoreward intrusions of Gulf Stream water onto the continental shelf off Jacksonville on 5
(solid line) and 10
(dashed line) November 1980. Note the filaments of mater stranded shoreward of the
intrusions. medrawn from Gulf Stream Frontal Analyses (GOES sea surface temperahue)
S. Baig, NOAA.]
Fig. 12. Association of Gymnodiuium breve cells with Gulf
Stream water in Onslow Bay between Cape Lookout and Cape Fear
during winter. ln a February 1991 cross-shelf transect of nine sta-
G. breve cells were found only at the two stations most close-
ly associated with the Gulf Stream meander. Water temperatures in
the Gulf Stream meander were -22°C; temperatures of the shelf
water did not exceed 14°C.
islands (Bane et #iI. 198 1; Yoder et al. 1985). These meanders
serve as nutrient pumps introducing new nitrogen from strata
beneath the Gulf’ Stream (Lee et al. 1991). After a meander
passes, parts of Ihe filament may remain on the shelf for as
long as a week before dispersing or rejoining the stream
(FRED Group 1989). The longevity (>I9 d) of the Gulf
Stream filament stranded on the continental shelf off North
Carolina in late fall 1987 was credited with sustaining this
unique G. breve bloom (Tester et al. 1991).
Significant qu#:stions remained in the aftermath of the
North Carolina bloom that caused the closure of major shell-
fish harvesting areas for an entire season and had an esti-
mated cost of $25 million (Tester and Fowler 1990). Samples
from ships of’ opr’ortunity were used to determine nonbloom,
background levels of G. breve for the northern and eastern
Gulf of Mexico, fhe Florida Strait, and the entire U.S. South
Atlantic Bight in:luding the Gulf Stream and western Sar-
gasso Sea (Fig. :!). G. breve has a continuous distribution
throughout this r;mge but in winter its occurrence in near-
shore waters of the U.S. South Atlantic Bight is closely as-
sociated with GLtlf Stream meanders overriding the shelf
(Tester et al. 1993). Perhaps the best example of this depen-
dency is from a cross-shelf sampling transect in Onslow Bay
(between Cape L’3okout and Cape Fear) in February 1991.
This transect bisected a Gulf Stream meander and G. breve
cells were found only in the meander and at its shoreward
edge (Fig. 12). Field studies of Rounsefell and Nelson
Tester and Steidinger
continue to operate on the bloom, cells concentrate in near-
shore waters where movement is governed by winds and
alongshore currents.
is found in low concentrations (< 1,000 cells
liter -I) in the Florida Current-Gulf Stream throughout the
year. During blooms cell concentrations of >2X lo4 cells li-
ter become entrained in the Florida Current and transported
to the Atlantic. Gulf Stream meanders are known to deliver
the cells into nearshore waters. Small-scale eddy features
like the Tortugas Gyre are very productive regions for phy-
toplankton and may prove to be so for G.
breve as
Strengthened transport through the Florida Strait at the dis-
sipation of the Tortugas Gyre suggests an explanation for the
infrequent, high cell counts in the Atlantic.
This conceptual model provides a framework to help de-
fine spatial and temporal scales, processes, timing, and link-
ages vital to hypothesis testing. It is essential that our un-
derstanding of’ the prerequisites for each bloom phase (e.g.
initiation, growth, maintenance, and dissipation) be as com-
plete as possible. We know little about factors that affect the
intensity OF G.
blooms. It is critical that surface cir-
culation be coupled with a depth component and the model
expanded to three dimensions. It is equally important to un-
derstand what factors contribute to the reintroduction of cells
into coastal waters-a major concern during prolonged
blooms. Increased access to archived wind and temperature
data combined with cell counts from “bloom logs” will aid
in retrospective analyses of blooms in greater detail. The
advent of online, real-time environmental data and the pros-
pect of new ocean color sensors may allow enough predic-
tive capability that bloom conditions can be detected early
enough to allow for focused research efforts and provide
reliable information to the public.
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... HAB events are implicitly spatially and temporally dependent, extending from a few square kilometers to thousands of square kilometers and persisting from a couple of weeks to a couple of months [12][13][14]. For a more effective detection, a combined spatial and temporal analysis is required. ...
... )Rrc(745) (12) Furthermore, the RDI results are shown in Figure 12. The HAB spatial range extracted by our EVA-based method was similar to the high RDI range, giving our method some credibility. ...
Full-text available
Long-term satellite observations have the ability to provide early warnings of harmful algal blooms (HABs). However, detecting HABs in optically complex coastal waters is somewhat challenging. In this article, we propose a two-step scheme, combining long short-term memory (LSTM) with extreme value analysis (EVA), for HAB detection. Essentially, the LSTM network builds a normal time series model on selected coordinate of long-term multisource satellite data. This model detects potential HAB dates by utilizing the LSTM predictive errors for an approximated Gaussian distribution. For each potential HAB date, the EVA approach then extracts the HAB distribution from the selected coordinate by considering the spatial correlation. A case study in Zhejiang coastal waters shows that our method exploits the advantages of both LSTM and EVA models, which not only has the strong prediction capability of LSTM for reducing HAB false alarm rate, but also achieves a dynamic HAB extraction through the EVA fitting.
... Heterocapsa sp. was even found to cause red tides in a eutrophic bay [48], and they often predominate under low temperatures and low light conditions [49]. Gymnodinium sp. can migrate in the water column [50], and its optimal temperature ranged from 7 to 24 °C [51], which explains the high abundance in the cold seasons that was observed in this study. ...
Full-text available
Syndiniales is a diverse parasitic group, increasingly gaining attention owing to its high taxonomic diversity in marine ecosystems and inhibitory effects on the dinoflagellate blooms. However, their seasonal dynamics, host interactions, and mechanisms of community assembly are largely unknown, particularly in eutrophic waters. Here, using 18S rRNA gene amplicon sequencing, we intended to elucidate the interactions between Syndiniales and microeukaryotes, as well as community assembly processes in a eutrophic bay. The results showed that Syndiniales group II was dominating throughout the year, with substantially higher abundance in the winter and spring, whereas Syndiniales group I was more abundant in the summer and autumn. Temperature and Dinoflagellata were the most important abiotic and biotic factors driving variations of the Syndiniales community, respectively. The assembly processes of microeukaryotes and Syndiniales were completely different, with the former being controlled by a balance between homogeneous selection and drift and the latter being solely governed by drift. Network analysis revealed that Syndiniales group II had the largest number of interactions with microeukaryotes, and they primarily associated with Dinoflagellata in the winter, while interactions with Chlorophyta and Bacillariophyta increased dramatically in summer and autumn. These findings provide significant insights in understanding the interactions and assembly processes of Syndiniales throughout the year, which is critical in revealing the roles of single-celled parasites in driving protist dynamics in eutrophic waters.
... Moreover, 'horizontal' exchanges between oceanic and coastal pelagic phytoplanktonic communities are usually observed. A flow from the ocean to coastal communities has been noticed for dinoflagellates especially [319,34]. Conversely, in many other bloomforming species, the shallower coastal areas might function as a reservoir for biodiversity in the ocean. ...
Full-text available
Phytoplankton communities, made of photosynthetic algae, can include up to hundreds of species requiring similar resources. Classical population dynamics models, however, often predict that the number of coexisting species cannot be much larger than the number of resources. Numerous explanations to this "paradox of the plankton" have been proposed, often based on the same hypotheses: interactions are competitive, population dynamics are based on a single life stage, corresponding to the organism floating in the water column (the pelagic stage), and these organisms are distributed homogeneously in space, all species being perfectly mixed in the environment. In this thesis, we build two independent models which enable us to relax these hypotheses.Firstly, we establish a community dynamics model with two life stages, involving a dormant one (a ‘seed’), which has a higher survival probability than the pelagic stage, especially in adverse environmental conditions. In this model, pelagic organisms can move between the ocean and the coast while dormant individuals remain in a coastal seed bank. The structure of interactions is inspired by field data, and comprises facilitation in addition to competition. The presence of a seek bank allows specialist species to survive in the community, and avoid the extinction of all species in harsh environmental conditions. Facilitation does not seem to promote coexistence.In the spatial section of the thesis, we present an individual-based model including hydrodynamic and demographic processes at the microscale. The replication of an existing single-species model in two dimensions allowed us to develop the numerical and analytical methods which serve as a foundation for a three-dimensional, multispecies model. In this model, birth and death events are modeled by a branching process, organisms are displaced by a random walk representing diffusion, and by a simplified model of turbulence. Parameter values are based on phytoplankton characteristics. We show that, for distances between individuals allowing interactions to happen, small organisms (nanophytoplankton) are mostly surrounded by individuals of the same species, which can favour coexistence, while larger species (microphytoplankton) are more mixed, which favours interspecific competition. We then discuss other potential mechanisms that could explain microphytoplankton diversity maintenance.
... The first source of nutrients comes from river outflows, which are also responsible for creating an ideal stratified area where dinoflagellates can out-compete other planktons [17]. The second source comes from the deep GoM and is brought to the continental shelf through upwelling processes [13,27,35]. Finally, large HABs also appear to be linked to aeolian supply of nutrients [32], but those constitute minor contributions and are insufficient to support large blooms [28]. ...
We investigate the influence of the Loop Current (LC) on the connectivity of the Gulf of Mexico, with a focus on the West Florida Shelf (WFS), using in situ trajectories from satellite-tracked drifting buoys in the Gulf of Mexico (GoM). We subset the dataset into two groups, Loop Current extended and retracted phases, that are used to construct two Markov Chains representing the distinct underlying dynamics during those two periods. The LC phases were found to impact and modify substantively the general connectivity of the GoM. Additionally, we highlight the presence of almost-invariant regions, where particles tend to remain for an extended period, on the WFS when the LC is extended. Those regions are bounded by a previously identified Cross-Shelf Transport Barrier and correspond to records of high-density areas of Karenia Brevis, a dinoflagellate responsible for red tides. Finally, we show that Markov Chain modeling could help forecast harmful algae blooms by using the devastating 2017 red tide event as a test case. The development phase on the WFS and its further propagation to Florida's east coast ecosystems is presented.
... The coastal transport of water and the vertical migration probably create optimum conditions for K. selliformis concentration near the coast. Similar mechanisms have been well described for K. brevis (Tester and Steidinger, 1997;Hetland and Campbell, 2007;Thyng et al., 2013). For example, K. brevis blooms typically initiate offshore in late summer or fall; regional hurricane and tropical storm activity typically peak during the same window . ...
In the fall of 2020, a long-lasting and massive harmful algal bloom (HAB) with extensive fields of yellow sea foam was observed in relatively cold waters (7–13 °C) off the coasts of the Kamchatka Peninsula, Russia. According to the estimates based on bio-optical parameters in satellite imagery, the Kamchatka bloom 2020 lasted for two months and covered a vast area of more than 300 × 100 km. An abundance of dead fish and invertebrates, including sea urchins, sea anemones, chitons, cephalopods, bivalves were found on shore during the bloom. Animals suffered almost 100% mortality within a depth range between 5 and 20 m. To identify the causative microalgal species, light and scanning electron microscopy, Raman spectroscopy, and molecular phylogenetic approaches were used. The HAB area was estimated by the spectral analysis of satellite-derived imagery. The causative organisms were unarmored dinoflagellates of Karenia species. Their density and biomass reached 100–620 cells·mL–1 and 1300–7700 mg·m–3, respectively, which accounted for 31–99% of the total cell density and 82–99% of the total phytoplankton biomass in late September to mid-October. The dominant species was Karenia selliformis, and the other co-occurring kareniacean species were K. cf. cristata, K. mikimotoi, K. papilionacea, K. longicanalis, and two unidentified morphotypes of Karenia spp. The molecular phylogeny inferred from LSU rDNA and ITS region showed that K. selliformis from Kamchatka in 2020 belonged to the cold-water group I and was identical to K. selliformis strains from Hokkaido, Japan, identified in 2021. This is the first HAB event caused by K. selliformis recorded from Russian coastal waters.
... Blooms of K. brevis are common along the coast of Texas and the southern Gulf coast of Florida, but less frequent along the Mississippi (MS), Alabama (AL), and Florida (FL) panhandle coasts (Tester andSteidinger 1997, Soto et al. 2018). More frequent K. brevis blooms on the southern Gulf coast of Florida may be due to higher nutrient inputs from the mainland (Brand andCompton 2007, Medina et al. 2022), whereas more frequent blooms on the Texas coast may be due to prolonged salinity increases (Tominack et al. 2020). ...
... Blooms of K. brevis are common along the coast of Texas and the southern Gulf coast of Florida, but less frequent along the Mississippi (MS), Alabama (AL), and Florida (FL) panhandle coasts (Tester andSteidinger 1997, Soto et al. 2018). More frequent K. brevis blooms on the southern Gulf coast of Florida may be due to higher nutrient inputs from the mainland (Brand andCompton 2007, Medina et al. 2022), whereas more frequent blooms on the Texas coast may be due to prolonged salinity increases (Tominack et al. 2020). ...
... Despite the success of K. brevis in the subtropical Gulf of Mexico, it has a more limited range than numerous other Karenia species and Kareniaceaens which coexist in the Gulf, though are rarely observed to bloom, despite causing devastating harmful algal blooms (HABs) around the world. Mechanisms of bloom initiation for K. brevis in the Gulf have been relatively well studied, and several contributing factors have been proposed (Tester and Steidinger 1997;Steidinger et al., 1998;Walsh et al., 2006;Hetland and Campbell 2007;Stumpf et al., 2008;Weisberg et al., 2014). Moreover, the traditional resting cyst life cycle phases common to other harmful dinoflagellate species have yet to be unequivocally identified in ✰ The Sanger nucleotide sequences reported in this article have been submitted to the NCBI GenBank under accession numbers MT162669-MT162673. ...
Karenia brevis, a neurotoxic dinoflagellate that produces brevetoxins, is endemic to the Gulf of Mexico and can grow at high irradiances typical of surface waters found there. To build upon a growing number of studies addressing high-light tolerance in K. brevis, specific photobiology and molecular mechanisms underlying this capacity were evaluated in culture. Since photosystem II (PSII) repair cycle activity can be crucial to high light tolerance in plants and algae, the present study assessed this capacity in K. brevis and characterized the ftsH-like genes which are fundamental to this process. Compared with cultures grown in low-light, cultures grown in high-light showed a 65-fold increase in PSII photoinactivation, a ∼50-fold increase in PSII repair, enhanced nonphotochemical quenching (NPQ), and depressed Fv/Fm. Repair rates were among the fastest reported in phytoplankton. Publicly available K. brevis transcriptomes (MMETSP) were queried for ftsH-like sequences and refined with additional sequencing from two K. brevis strains. The genes were phylogenetically related to haptophyte orthologs, implicating acquisition during tertiary endosymbiosis. RT-qPCR of three of the four ftsH-like homologs revealed that poly-A tails predominated in all homologs, and that the most highly expressed homolog had a 5′ splice leader and amino-acid motifs characteristic of chloroplast targeting, indicating nuclear encoding for this plastid-targeted gene. High-light cultures showed a ∼1.5-fold upregulation in mRNA expression of the thylakoid-associated genes. Overall, in conjunction with NPQ mechanisms, rapid PSII repair mediated by a haptophyte-derived ftsH prevents chronic photoinhibition in K. brevis. Our findings continue to build the case that high-light photobiology—supported by the acquisition and maintenance of tertiary endosymbiotic genes—is critical to the success of K. brevis in the Gulf of Mexico.
... Weise et al. (2002) reported windy conditions could prevent the development of Alexandrium tamarense blooms or cause their dissipation in the St. Lawrence Estuary [17]. The mixing or disruption of a water mass, especially in combination with declining water temperatures, was also considered important in the dissipation of Karenia brevis blooms in the Gulf of Mexico [18,19]. The low salinity water from A bloom event occurred in the Pearl River Estuary during the August of 2011. ...
Full-text available
Dinoflagellates is one dominant group in coastal marine phytoplankton communities and, on occasion, form blooms in estuaries and coastal ecosystems. While relationships between dinoflagellate bloom dynamics and nutrients are well-studied, information regarding bloom dis-sipation in estuaries is limited. We studied the dissipation of dinoflagellate Polykrikos geminatum blooms in the Pearl River Estuary, South China Sea, during August of 2011 using ecological, molecular , and satellite remote sensing data. We found that the dinoflagellate bloom was associated with water temperatures of 29.2-31 °C, salinities ranging 16.4-20, and ambient water nutrient concentrations that were not limited. The abundance of the ciliate Euplotes rariseta, which feeds on P. geminatum cell debris and bacteria, functions as an indicator species of P. geminatum bloom dis-sipation. In situ and satellite data indicate that bloom water masses were transferred from the central to inner estuary near Shenzhen Bay, driven by continuous, strong southerly winds; at which point in time, P. geminatum blooms dissipated to a high-salinity area near the estuary mouth driven by northerly winds and freshwater discharge, whereupon the blooms rapidly vanished. A low tolerance to low or high salinities resulted in P. geminatum bloom demise in the Pearl River Estuary. We propose that interactions among salinity, wind, and freshwater incursion result in P. geminatum bloom dissipation in the Pearl River Estuary.
Red tide events, caused by a toxin producing dinoflagellate, Karenia brevis, occur annually in Florida and Texas. These events lead to health risks for both humans and wildlife that utilize coastal environments. Brevetoxins, potent lipophilic neurotoxins produced by K. brevis, modulate immune responses in laboratory studies with model organisms and in the natural environment in both humans and wildlife. Studies show that brevetoxins activate immune cells, stimulate production of gamma-globulins, cytokines, and neutrophils, modulate lysozyme activity, induce apoptosis, and modulate lymphocyte proliferation in marine species. The objective of this review was to summarize brevetoxin-induced immunotoxicity in marine animals based on available peer-reviewed literature about K. brevis blooms and associated health concerns and propose putative toxicity pathways. This review identifies knowledge gaps within current brevetoxin induced immunotoxicity research, including assessing the long-term impacts of brevetoxin exposure, elucidating the mechanistic linkages between brevetoxins and immune cells, and evaluating repeated and chronic versus acute brevetoxin exposure implications on overall organismal health. The putative immunotoxicity pathways based on evidence from brevetoxin-exposure in marine fauna described in this review represent a useful tool and resource for researchers, wildlife managers, and policy makers. This review and putative immunotoxicity pathways will inform decisions regarding the risks of algal blooms, as it pertains to marine animal health. Marine animal veterinarians may also use the putative immunotoxicity pathways as a reference to guide research on supportive care, treatment, and preventatives.
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An intrusion of loop current water up DeSoto Canyon and onto the West Florida continental shelf to within 8 km of the shore occurred in February 1977. Boat, aircraft, and satellite data collected in the area for another purpose were used to estimate the space and time scales of the intrusion and the ultimate fate of the intruded waters. The duration of the event was 18 days. Oceanic waters advanced across the shelf at speeds of 20 cm s−1. At maximum intrusion, 6650 km2 of shelf were affected. Approximately half the intruded water receded off the shelf, and half appears to have been modified in situ.
Simultaneously measured Eulerian currents and spatially extensive subsurface temperatures have provided a time series of eight synoptic, three-dimensional views of the Gulf Stream frontal zone along the Carolina continental margin. Two large-amplitude meanders were observed to progress through the study area between Charleston and Cape Hatteras during February 1979. Each meander had a vertically coherent, skewed wave-like subsurface structure. The Eulerian velocity and temperature signatures produced by the meanders at the 250-m level over the 390-m isobath reflect this skewness. At a particular instrument, the in-phase increases in temperature and downstream velocity associated with an approaching meander crest occurred during a longer time interval than did the more rapid decreases in these quantities following the crest's passage. Typically, the downstream velocity component at this level fluctuated from about -20 cm s-1 to near 100 cm s-1, while the cross-stream component varied approximately ±25 cm s-1 about a near-zero mean. For a particular meander, the maximum in the offshore velocity component led the downstream maximum in time in a manner typical of progressive wave motions; however, the lead time was always less than one quarter of a meander period implying that u and υ were not in quadrature, as would have been the case for stable waves. The two meanders were observed downstream of the area off Charleston where a seaward deflection of the stream is often found. Subsurface temperature data from February 10, 1979, show that on that date the degree of deflection was greatest near the surface, and that almost no deflection existed within the deeper reaches of the water column. According to later data, the deflection decreased as the meanders progressed alongshore away from the area, suggesting that the vertical structure of the deflection observed on the tenth may have been associated with the late stages of a meander passage. Filaments of warm Gulf Stream water extended southwestward `behind' the crests of the two meanders. The filaments were relatively shallow features, extending from the surface to a depth of a few tens of meters. They were oriented essentially parallel to the bottom contours over the outer shelf and upper slope, and were separated from the main body of the Gulf Stream by cool water. The presence of the cool water between the stream and the filaments at the surface was due to upwelling of water from deep within or below the main stream. Peaks in the time series of vorticity components indicate that maximum cyclonic relative vorticity occurred behind the meander crests, in the leading portion of the trough near where a warm filament joined a meander crest. The meanders may have been initiated upstream of our study area, and then `amplified' by the deflection process off Charleston. Energy flux calculations for the region off Onslow Bay indicate that meander kinetic energy was being converted to mean energy there. It seems likely that the deflection produces meander growth within the 100 km or so immediately downstream of Charleston, to subsequently have the meander energy reconverted to mean energy farther downstream.