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Factors Affecting the Distribution of Pyrodinium Bahamense Var

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Factors Affecting the Distribution of Pyrodinium Bahamense Var MARINE ECOLOGY PROGRESS SERIES Vol. 322: 99–115, 2006 Published September 20 Mar Ecol Prog Ser Factors affecting the distribution of Pyrodinium bahamense var. bahamense in coastal waters of Florida E. J. Phlips*, S. Badylak, E. Bledsoe, M. Cichra Dept. Fisheries and Aquatic Sciences, University of Florida, 7922 N.W. 71st Street, Gainesville, Florida 32653, USA ABSTRACT: Over the last half of the 20th century Pyrodinium bahamense var. bahamense has been observed in a variety of locations in the western North Atlantic. Recent evidence of the toxin- producing capacity of this variety of P. bahamense has heightened interest in its habitat requirements and preferences. The objective of this study was to examine the environmental factors that relate to the spatial and temporal patterns of the distribution and abundance of P. bahamense var. bahamense. Based on the results of this study we view the factors as operating in one or more ways: (1) eco- physiological limitations for survival and successful reproductive cycle, (2) environmental regulation of growth and standing crop, and (3) competitive advantages in relation to other species. The focus of the study was the Florida peninsula, but information from other environments in the tropical Atlantic and Gulf of Mexico was included in the interpretation of the results. In terms of physiological limitations, 20°C appears to be the lower temperature limit for a significant presence of P. bahamense var. bahamense, and the salinity tolerance ranged from 10 to 45. The bloom potential of P. bahamense var. bahamense was most closely associated with shallow ecosystems with long water residence times, and peak biomass levels were correlated to nutrient concentrations in regions of high abundance. The ability of P. bahamense var. bahamense to compete effectively for habitat with other euryhaline warm-water phytoplankton is viewed in terms of existing theories on succession and competition, including Margalef’s Mandala, Reynolds’ Intaglio and C-S-R life-form strategies proposed by Smayda & Reynolds. KEY WORDS: Phytoplankton · Succession · Tropical Atlantic Ocean · Gulf of Mexico · Dinoflagellate · Harmful algae · Saxitoxin Resale or republication not permitted without written consent of the publisher INTRODUCTION light emissions attributable to the dinoflagellate. In the early accounts of P. bahamense var. bahamense, The prominence of the toxic bioluminescent dino- no distinct varietal status was assigned to the Atlantic flagellate Pyrodinium bahamense var. compressum in form, but subsequent morphological studies suggest the tropical Indo-Pacific region has been extensively that such a distinction is warranted (Steidinger et al. described (Maclean 1989, Shumway 1990, Hallegraeff 1980, Badylak et al. 2004), although not universally 1993, Smayda 1997, Anton et al. 2000, Azanza & Tay- accepted (Balech 1985). Over the last half of the 20th lor 2001); however, the distribution of its Atlantic century P. bahamense var. bahamense has been Ocean counterpart, P. bahamense var. bahamense, is observed in a variety of additional locations in the less well defined. One of the earliest reports of P. western North Atlantic, including Oyster Bay in bahamense in the North Atlantic came from the Jamaica (Seliger et al. 1970), and Florida Bay (Phlips Bahamas by Plate (1906). Later, Margalef (1961) & Badylak 1996), Indian River Lagoon (Badylak & reported blooms of P. bahamense in Phosphorescent Phlips 2004), and Tampa Bay (Steidinger et al. 1980, Bay, an aquatic habitat in Puerto Rico named for the Badylak et al. 2006) in Florida. *Email: [email protected] © Inter-Research 2006 · www.int-res.com 100 Mar Ecol Prog Ser 322: 99–115, 2006 One of the most noteworthy features of Pyrodinium are discussed within the context of current hypotheses bahamense is the ability of the Indo-Pacific form, regarding the factors that favor blooms of dino- P. bahamense var. compressum, to produce saxitoxin, a flagellates (Margalef et al. 1979, Reynolds & Smayda powerful neurotoxin responsible for widespread 1998, Smayda & Reynolds 2001), including nutrient human mortalities in the Indo-Pacific region (Maclean status, hydrologic conditions, and biological inter- 1989, Azanza & Taylor 2001). Until recently, it had been actions. As suggested by Reynolds and Smayda (Rey- assumed that the Atlantic form of P. bahamense did not nolds & Smayda 1998, Smayda & Reynolds 2001), ‘life produce the neurotoxins associated with P. bahamense form properties, habitat preferences and stochastic var. compressum (Steidinger et al. 1980, Azanza & selective processes’ are more important in defining the Taylor 2001). However, recent studies of saxitoxin character of succession and the magnitude of blooms bioaccumulation in pufferfish from the Indian River La- than broad phylogenetic associations. goon in Florida led to the discovery that local strains of P. bahamense (var. bahamense) can produce the toxin (Landsberg et al. 2002). This discovery has heightened MATERIALS AND METHODS interest in the factors that regulate the presence and abundance of P. bahamense var. bahamense. Study site descriptions. In an effort to describe the Another recent observation about the Atlantic form distribution and prominence of Pyrodinium bahamen- of Pyrodinium bahamense var. bahamense is its ability se var. bahamense within Florida, phytoplankton com- to form blooms under low-salinity conditions (i.e. salin- position data gathered by the authors since 1993 and ities <25) (Phlips et al. 2004a). Such conditions are information from existing literature were both used in found in many lagoons and estuaries in Florida. It had this study. In the former category, data were available been previously hypothesized that blooms of P. baha- for 20 sampling sites along the east coast of Florida, mense var. bahamense were restricted to regions of 18 sites in Florida Bay at the southern tip of Florida, higher salinities (i.e. salinities >30) (Wall & Dale 1969), and 34 sites along the western coast of the Florida much like the prominent toxic dinoflagellates Karenia peninsula, focusing on Tampa Bay and the Suwannee breve and Gamberodiscus toxicus. This observation River estuary (Fig. 1). expands the potential realm of influence for P. baha- The northernmost sampling sites along the east coast mense var. bahamense. of Florida were located in the mesohaline portion of the The objective of the present study was to examine St. Johns River. The lower reach of the St. Johns River the environmental factors that relate to the spatial and extends 175 km, from Lake George to the Atlantic temporal patterns of the distribution and abundance of Pyrodinium bahamense var. bahamense. Based on the results of St. Johns River (3 sites) this study we view the factors as operat- SSt.t. AAugustineAugustinugustine Tolomato/Gauna to ing in one or more ways: (1) ecophy- PalmPalm CoasCCoastoast Matanzas River (6 sites) siological limitations for survival and DaytonDDaytonaaytona Halifax River (3 sites) successful reproductive cycle, such as Mosquito Lagoon (1 site) Suwannee River temperature and salinity tolerances; (2) Perdido Bay Estuary (34 sites) TTitusvilleTitusvillitusville environmental regulation of growth and EauEau GalliGGallieallie Indian River standing crop, such as nutrient availabil- MMelbourneMelbournelbourne Lagoon SebastianSSebastiaebastian ity; and (3) competitive advantages in Choctawhatchee Bay (7 sites) Tampa Bay VVeroVerero relation to other species, such as growth Escambia/Pensacola Bay (3 sites) rate potential. The focus of the study was the Florida peninsula, but information from other environments in the tropical Miami Atlantic and Gulf of Mexico was in- cluded in the interpretation of the results. The study was aided by the large variety of environmentally distinct coas- Florida Bay (18 sites) tal habitats encountered in Florida. Florida is also known as a biogeographi- cal transition zone from the tropical Fig. 1. Locations of sampling sites on the peninsula of Florida. In addition to sites examined in the current study, literature accounts of phytoplankton stud- habitats of the Florida Keys to the warm ies were also considered from 3 locations in the panhandle of Florida. Regions temperate regions of the Georgia and where Pyrodinium var. bahamense was not observed or reported by other Carolina coasts. The results of this study researchers are shown in italics Phlips et al.: Pyrodinium bahamense distribution 101 Ocean. Due to the very shallow slope of the river basin, regions of the bay and the Atlantic Ocean and Gulf of the influence of oceanic water can extend up to 75 km Mexico. The locations of the 18 sites were selected to upstream of the outflow. Much of the river is shallow, be representative of the major sub-basins within the i.e. <4 m in depth, with the exception of the dredged lagoon (Phlips et al. 1996, 1999). Water was collected navigational channels. Phytoplankton composition monthly from 1993 to 1997. data were available for 3 sampling sites within the Along the west coast of Florida 2 major regions were mesohaline portion of the river on a monthly basis for a included in the sampling effort, Tampa Bay and the 12 yr period from 1993 through 2005. The 5 sites were Suwannee River estuary. Tampa Bay is Florida’s lar- located from 4 to 70 km from the outflow. gest estuary, with a surface area of 1031 km2. The estu- South of the St. Johns River, the sampling sites along ary has a mean depth of 3.7 m (Schmidt & Luther 2002). the east coast of Florida were mostly located in the Water samples were collected from 3 sampling sites in lagoonal habitats between the barrier islands and the bay, 1 near the mouth of the bay, 1 in the center of mainland of Florida. The Intra-Coastal Waterway the bay, and 1 in the northern basin of the bay, known extends for almost the entire length of the east coast of as Old Tampa Bay. The latter sampling site is charac- Florida connecting the various lagoons. All of the la- terized by long residence times caused by the rela- goons are characterized by shallow mean depth, i.e. tively small freshwater inflows to the northern basin generally <3 m, except in dredged navigational chan- and the physical restriction to tidal water exchange nels.
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