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CONTENTS
Introduction
Materials and Methods
Results
Discussion
In Memoriam
Acknowledgments
Literature Cited

Northeast Fisheries Science Center Reference Document 03-13

Distribution of the Brown Tide Picoplankter Aureococcus anophagefferens in Western New York Bight Coastal Waters

John B. Mahoney1, Dorothy Jeffress2, Christine Zetlin1, Paul S. Olsen3, Helen Grebe4, and Jasen Brooks1
1National Marine Fisheries Service, James J. Howard Marine Sciences Laboratory, 74 Magruder Road, Highlands, NJ 07732; 2National Marine Fisheries Service, Milford Laboratory, 212 Rogers Avenue, Milford, CT 06460; 3New Jersey Department of Environmental Protection, Bureau of Freshwater and Biological Monitoring, P. O. Box 427, 35 Arctic Parkway, Trenton, NJ 08625; 4U. S. Environmental Protection Agency, 2890 Woodbridge Ave., Edison, NJ 08837

Web version posted September 25, 2003

Citation: Mahoney, J.B.; Jeffress, D.; Zetlin, C.; Olsen, P.S.; Grebe, H.; Brooks, J. 2003. Distribution of the brown tide picoplankter Aureococcus anophagefferens in western New York Bight coastal waters. Northeast Fish. Sci. Cent. Ref. Doc. 03-13; 23 p.

Information Quality Act Compliance: In accordance with section 515 of Public Law 106-554, the Northeast Fisheries Science Center completed both technical and policy reviews for this report. These predissemination reviews are on file at the NEFSC Editorial Office.

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ABSTRACT

Incidence of the picoplankter Aureococcus anophagefferens in eastern Long Island, NY, has been thoroughly documented since its blooms, or "brown tides", began there in1985. In contrast, definitive information on incidence of A. anophagefferens in the western or New Jersey side of the New York Bight lagged considerably. None was available until surveys along the northeast U. S. coast in 1988 and 1990 by other investigators detected the species in New Jersey bays and ocean coastal waters from the Hudson-Raritan estuary south to Great Bay (approximately central on the New Jersey coast). Confirmation of an A. anophagefferens bloom in New Jersey, this in the Barnegat Bay-Little Egg Harbor estuarine system, was delayed until 1995 although earlier episodes were suspected. To obtain more comprehensive and current information on A. anophagefferens distribution and potential for its blooms in the western Bight, we surveyed for it in coastal waters from Delaware Bay to the Hudson-Raritan estuary, and in western Long Island (Nassau County, NY) south shore locales, during 1997 to 2001. Results showed persistence of the species in New Jersey locales where found by the 1988,1990 surveys, and expansion of its range southward in New Jersey coastal waters since 1990. Year-to-year difference in incidence in waters south of Great Bay was noted, with much greater incidence in1999, a year when it bloomed primarily in the Barnegat Bay-Little Egg Harbor system and Great Bay, than in1998, a non-bloom year. Cell numbers in 1999 in New Jersey southern estuarine waters were below reported detrimental level (3.5 x 104 cells ml-1 ) at most sites surveyed, but at three sites to approximately six km south of Great Bay concentrations ranged 2-2.8 x 106 cells ml-1. This was the first confirmed bloom occurrence south of Great Bay. Although having a history of intense blooms of other phytoplankton species, the Hudson-Raritan estuary does not appear to be a system where brown tide might be expected.

KEY WORDS: Harmful algal blooms, brown tide distribution, New York Bight, Aureococcus anophagefferens.


INTRODUCTION

Intense blooms of a previously unidentified picoplankter in New York Bight coastal waters were reported first in the mid-1980's in eastern Long Island bays, including the eastern-most Gardiners Bay-Peconic Bay system, and bays along the southeastern shore including Shinnecock Bay, Moriches Bay, and Great South Bay (Nuzzi, 1995; Bricelj and Lonsdale, 1997). The causative picoplankter was described and named Aureococcus anophagefferens by Sieburth et al. (1988). Early recognition of severe detriment of the eastern Long Island blooms, or "brown tides", to bay scallop, Argopecten irradians, and other components of the biota (Bricelj and Kuenster, 1989) resulted in considerable timely attention to these blooms. Examination of brown tide as a regional phenomenon has been inadequate, however, despite a documented 1985 A. anophagefferens bloom in Narragansett Bay, RI (Smayda and Villareal, 1989). A. anophagefferens incidence information for the western or New Jersey side of the New York Bight was lacking completely until Anderson et al. (1993) provided limited data. A 1995 brown tide episode was the first confirmed in the western Bight (Nuzzi et al., 1996); such confirmation may have been delayed for a decade. New Jersey possibly experienced A. anophagefferens blooms in 1985-1987 but the dominant picoplankter was not identified definitively (Olsen, 1989). A. anophagefferens could not be distinguished reliably from similar picoplankters using light microscopy. Means for positive identification of the species was not available to monitoring agencies until Anderson et al. (1989) developed an immunofluorescence protocol for this.

The goal of the Anderson et al. (1993) surveys for A. anophagefferens along the northeast U. S. coast in1988 and 1990 was to determine its distribution beyond the eastern Long Island embayments bloom loci. The southerly limit of the 1988 survey was Barnegat Bay at Manahawkin, New Jersey; their 1990 survey extended to Chesapeake Bay. They found the species in New Jersey bays and ocean coastal waters from the Hudson-Raritan estuary south to the Barnegat Bay-Great Bay sector (approximately central on the New Jersey coast). In 1988, New Jersey samples were collected on September 20, well past the usual time for primary bloom development and maxima (May-June). Nevertheless, alarming levels of A. anophagefferens (3.5-14.1 x 104 cells ml-1) were detected in southern Barnegat Bay. Brownish water discoloration characteristic of an A. anophagefferens bloom was not observed in the bay in 1988 (Olsen, unpublished data). Such discoloration is evident when A. anophagefferens cell numbers are > 2.0 x 105 ml-1 (Nuzzi, personal communication). If a low level bloom occurred there in1988 it likely was masked, because total picoplankton concentrations (likely dominated by Nannochloris atomus) approximated 2.0 x 106 cells ml-1 and A. anophagefferens when assessed comprised only ~ 7.5% of the picoplankton (Olsen, 1989). The Anderson et al.1990 survey sampling again was post-bloom maximum time, August 29, and when water temperature likely was unfavorably high for A. anophagefferens (i.e. >26°C). They detected the species in low numbers (highest, 216 cells ml-1) in several Barnegat Bay locales and in Great Bay, but not between Great Bay and Chesapeake Bay.

Anderson et al. (1993) concluded that the widespread distribution of A. anophagefferens they found in waters far from the eastern Long Island, New York, population "center" suggests that numerous areas have the potential for destructive brown tides. They recommended continued monitoring for the species. We conducted surveys for A. anophagefferens in 1997 to 2001, from Delaware Bay to western Long Island, to update and expand information on distribution of the species in this area, and to identify additional locales where its blooms might be expected. We report the initial A. anophagefferens distribution information for western Long Island; Nuzzi (personal communication) had advised there was a dearth of such information. This report excludes A. anophagefferens incidence information for the brown tide-prone Barnegat Bay-Little Egg Harbor system (Mahoney et al., 1999) which will be reported separately.


MATERIALS AND METHODS

The New York Bight is bounded by easternmost Long Island on the north and southernmost New Jersey on the west. Surveys for A. anophagefferens were conducted on an irregular basis in coastal ocean waters, embayments, and intracoastal waters of New Jersey and western Long Island during1997-2001. The initial survey, in July 1998, was of bay and intracoastal waterway sites from Delaware Bay to the Hudson-Raritan estuary, excluding the Barnegat Bay-Little Egg Harbor system. Survey of such sites from Delaware Bay to Great Bay was repeated on September 30, 1998, June 14-16, 1999, and May 31, 2000. Samples were collected occasionally at nine New Jersey intracoastal and bay sites and thirteen New Jersey coastal ocean sites between 1995 and 2000. A total of 14 western Long Island estuarine and coastal ocean sites were sampled in June 2001. Survey sites are shown in Figure 1 (survey area with sites numbered), and Figure 1A, Figure 1B, Figure 1C, and Figure 1D (survey area regions with sites numbered and named). Combined letter/number designations in parentheses denote standard U. S. Environmental Protection Agency (EPA) sampling stations. Figure site numbers correspond to site numbers in Tables 1, 2 and 3.

Bay water samples primarily were collected from docks along shore with a Niskin bottle at ~0.5 m depth. Offshore bay samples were collected by helicopter (EPA, Region II) with a Kemmerer bottle at ~1.0 m depth. Helicopter collections (same depth and method) also provided coastal ocean samples, from ~0.4 km to ~1.6 km from shore. Water salinity and temperature measurements were made by personnel of various agencies and by different means. Salinity was measured by Yellow Springs Instrument Co. meter or refractometer, and is expressed as practical salinity units (PSU), equivalent to parts per thousand (0/00). Water temperature measurements were made variously by meter (Orion Model 265; Hanna Model HI 9060) and thermometer.

Sample handling, preservation and immunofluorescence protocol for A. anophagefferens identification and enumeration basically were that of Anderson et al. (1989, 1993). Bureau of Marine Resources, Suffolk County, NY, Department of Health Services (SCDHS) enumerated A. anophagefferens in 1995 and some 1997 samples. Otherwise, A. anophagefferens enumerations were done at the James J. Howard Marine Sciences Laboratory (HL). SCDHS provided initial training and continuing advice in the immunofluorescence protocol to HL personnel. A. anophagefferens identification and enumeration was confirmed periodically between the two labs. A. anophagefferens population levels at collection sites are provided in Tables 1, 2 and 3, along with associated water temperature and salinity when available. Sample immunofluorescence preparations were scanned for presence of A. anophagefferens prior to enumeration. When the species was seen during the scan but not during enumeration its presence "P" was tabulated. Bricelj et al. (2001) reported that A. anophagefferens concentrations as low as 3.5 x 104 cells ml-1 reduced feeding of juvenile hard clam, Mercenaria mercenaria. This level will be referred to in regard to relevance of A. anophagefferens population levels detected.


RESULTS

New Jersey Coastal Ocean Sites

Atlantic Ocean coastal waters, from off Cape May Point, southernmost on the New Jersey ocean shore, to the northern ocean shore off Monmouth Beach (Figure 1), were sampled irregularly during May through August, 1997 to 2000. One site also was sampled in 1995. Sites included some ~1.6 km from shore as well as the majority which were ~0.4 km from shore. Results are shown in Table 1. A. anophagefferens was found at least part of the time at all sites; levels ranged to 4.6 x 103 cells ml-1 (site 3, off Cape May Point, Figure 1A) but predominantly were <300 cells ml-1 over the several years of the study. Highest levels of the species were found in southern coast and mid-coast sites (sites 3, 39, 45) in1999. This was concurrent with or shortly after an intense bloom in the Barnegat Bay-Little Egg Harbor system, Great Bay, and some contiguous bays to the south (Figure 1B). Sites sampled in multiple years (sites 3, 49, 54), with the exception of site 3 in1995, had presence of A. anophagefferens each year. The greatest year-to-year change in abundance was at site 3. Sites 49 and 54, sampled multiple times during particular years, had varied same-year A. anophagefferens presence, i.e., from 0 to ~300 cells ml-1. Comparison of geometric means of cell levels (multiple enumerations for individual sites were averaged) at southern shore sites (Cape May to Atlantic City - sites 3, 5, 12, 13, 17, 19) and northern shore sites (Manasquan to Monmouth Beach - sites 49, 50, 54) shows comparable levels, 149 vs.151cells ml-1.

New Jersey Bay and Intracoastal Sites

Survey for A. anophagefferens in New Jersey bay or intracoastal waters, excluding the Barnegat Bay-Little Egg Harbor system and Great Bay (excepting southern shore) (Figure 1B), primarily was done irregularly during May-September, 1997-2000. Results are shown in Table 2. The species was present from Delaware Bay to the Hudson-Raritan estuary at most sites, most of the time. It was detected more in the southern region, from Delaware Bay to the southern shore of Great Bay (Figure 1A), than in the northern region, from Shark River to the Hudson-Raritan estuary (Figure 1C). A. anophagefferens was found at 16 of 20 sites in the southern region (sites 1-26), whereas in the northern region (sites 51-71) the species was found at only eight (sites 51, 55, 62, 63, 64, 66, 69, 71) of 20 sites. However, 17 of the latter were sampled only in1998, a non-bloom year, but15 of 20 southern sites were sampled in one or more bloom years, i.e., 1995, 1997, 1999, 2000. Instances of non-detection in the southern region during the course of the surveys were associated with sites sampled only in1998 (sites 8, 12, 20, 22). In 1998, the species was found at only six (sites 1, 6, 14, 15, 23, 24) of 15 sites sampled in the southern region, although at site15, Great Egg Inlet, it was found in three samplings over a month. In contrast, in 1999 it was present at all southern region sites. Cell levels at seven of 10 southern region sites (sites 6, 9, 10, 14, 15, 18, 24) sampled in 1999 and at least one other bloom year, e.g. 2000, were highest in 1999; at two sites (sites 4,16) levels were higher in 2000, and at one site (site 5) levels for 1999 and 2000 were low and comparable. Sites in the Hudson-Raritan estuary (Figure 1C, Table 2, sites 62 to 71) primarily were sampled in May through August, 1997, 1998. We confirm presence of the species in Sandy Hook Bay (sites 62-64, 66, 69) reported by Anderson et al.(1993) and also report its presence in Raritan Bay (site 71).

Data for Oyster Creek at the south shore of Great Bay (site 27) are included in Table 2, although this bay is considered in the central region. Shown is that A. anophagefferens was undetected there in a single September 1998 sample, a high level (1.7 x 106 cells ml-1) was present in June 1999, and low levels < 1000 cells ml-1 were present in June-August 2000.

A. anophagefferens population levels in1999 were below reported minimum detrimental level (3.5 x 104 cells ml-1 ) at most sites, but three sites to approximately six km south of Great Bay, i.e., Reed Bay, Perch Cove and Obes Thorofare, Brigantine (Figure 1A; Table 2, sites 24, 25, 26) had concentrations ranging 2-2.8 x 106 cells ml-1.

Western Long Island South Shore Sites

A. anophageffens was present at all fourteen western Long Island south shore sites (Table 3, sites 72-85) sampled during the first three weeks of June, 2001. At thirteen of these sites, sampled multiple times, cell levels increased slightly during the month. Highest cell levels (~1000-2000 cells ml-1) were found in coastal ocean samples from the vicinity of Jones Inlet (sites 3, 74, 75); considerably lower levels were found in this Inlet (site 78).

Salinity and Temperature Conditions

Encompassing open ocean and estuarine locations from Portsmouth, New Hampshire to the Chesapeake Bay, Anderson et al. (1993) detected A. anophageffens in waters having salinities ranging from18-32 PSU. In our western New York Bight surveys we detected it in a similar salinity range:18.5-34 PSU; the temperatures of waters we found it in ranged 14.1-28.3°C.


DISCUSSION

Prior to the surveys we report, the only available A. anophagefferens incidence information for the western New York Bight was that of Anderson et al. (1993) who provided data from single samplings of five sites in September, 1988 and eight sites in August, 1990. Our results are insufficiently comprehensive to support conclusions about A. anophagefferens population dynamics in western New York Bight waters, but are a substantial increase in incidence information. It is clear now that the species is distributed along the whole coast of New Jersey, in coastal ocean waters, and especially in certain embayments. It appears to be firmly established in some embayments as a constituent of the phytoplankton community. A. anophagefferens does not flourish unless environmental conditions are suitable. Comparison of 1998 and 1999 incidences shows that distribution of the species, and suitability of a body of water to support high concentrations of the species, are best gauged when regional environmental conditions are favorable. Our results also show that single assessments of incidence can be unreliable, and multiple assessments through a growing season are necessary.

In New Jersey coastal ocean waters A. anophagefferens evidenced consistent presence in relatively low abundance (usually <300 cells ml-1). Although detected at ~5000 cells ml-1 at a coastal ocean site in June, 1999 -- when it bloomed intensely in the Barnegat Bay-Little Egg Harbor system, Great Bay, and some contiguous bays to the south -- it was never found in high abundance in ocean waters. Difference in mean levels between New Jersey southern and northern coastal ocean waters was not apparent. Comparison of limited data available from a decade earlier suggests little change in levels of the species; Anderson et al.(1993) reported levels of 49 and 243 cells ml-1, respectively, at two New Jersey coastal ocean sites in their 1990 survey. Presence of A. anophagefferens along the northern half of the New Jersey ocean shore detected by Anderson et al. (1993) in their 1990 survey, is confirmed to the year 2000. Anderson et al. (1993) did not detect the species south of Great Bay, NJ. The present study, therefore, extends its known range in western New York Bight coastal ocean waters south to Delaware Bay. It is noted that in 1998, a non-bloom year, all five coastal sites south of Atlantic City sampled had presence of A. anophagefferens (range 36-118 cells ml-1), whereas only one of six intracoastal sites in this area had it present, and this below detection level for enumeration.

Sampling the southern region in bloom years (1995, 1997, 1999, 2000) and sampling the northern region primarily in 1998, a non-bloom year, likely skewed results but, nevertheless, the data do suggest greater incidence in the southern region. Supporting this is that there are more bodies of water potentially of high suitability for the species in the southern coastal region than in the northern coastal region. Central on the New Jersey coast, the Barnegat Bay-Little Egg Harbor estuarine system, New Jersey's principal barrier island system, has a history of brown tide occurrence. South of this system, a similar complex of bays or "sounds"connected by intracoastal channels (including the Intracoastal Waterway) extends along the New Jersey coast. In contrast, to the north, the Metedeconk River, Manasquan River and Shark River are the only sizeable estuarine waters between Barnegat Bay and the Hudson-Raritan estuarine system. Generally higher incidence of A. anophagefferens in the southern region in1999, relative to other bloom years, suggests especially favorable conditions that year. Higher incidence and occurrence of its blooms in certain New Jersey embayments follows a similar pattern of its incidence and blooms on Long Island. Paralleling its distribution in abundance on Long Island being limited to embayments and barrier island systems in the eastern region, at distance from the Hudson-Raritan estuary and adjacent waters, greatest incidence and abundance in New Jersey was found in the central and southern coast regions.

During the June1999 A. anophagefferens bloom, high levels (2-2.8 x 106 cells ml-1) were detected, to approximately six km south of Great Bay, in Reed Bay, Perch Cove and Obes Thorofare, Brigantine. This was the first confirmed bloom occurrence south of Great Bay. These cell levels are comparable to the maximum reported for eastern Long Island (Bricelj and Lonsdale, 1997). Because Absecon Bay is directly contiguous with Reed Bay, the bloom likely was present there also, but Absecon Bay unfortunately was not sampled at the time. A June 1999 level of 2.5 x 104 cells ml-1 detected in Beach Thorofare at Atlantic City (Table 2, site 21), about 13 km south of Great Bay, is further evidence of southward occurrence of the bloom. This population was below reported detrimental level when sampled. However, A. anophagefferens concentration can approximately double in a day (Dzurica et al., 1989). Beach Thorofare is contiguous with Absecon Bay on its south side; the elevated level detected in the Thorofare supports suspicion that the species bloomed in Absecon Bay. The comparably high A. anophagefferens level (1.7 x 106 cells ml-1) at Oyster Creek at the south shore of Great Bay at this time suggests continuity of the bloom in the Barnegat Bay-Little Egg Harbor system with bloom occurrence to the south.

The Hudson-Raritan estuary, where high primary productivity is characteristic (O'Reilly et al., 1976), and where intense blooms of various phytoplankton species have recurred over the last several decades (Olsen and Mahoney, 2001), does not appear to be a system where brown tide might be expected. A. anophagefferens has been present in the estuary for at least nine years (Anderson et al.,1993), and was detected there only in relatively low numbers in years when it was blooming in the Barnegat Bay-Little Egg Harbor system. Its long term-presence but apparent inability to bloom in the Hudson-Raritan estuary suggests water quality unfavorable to the species may be a factor. Steele et al. (1989) found A. anophagefferens to be among the most sensitive species of marine flora and fauna tested to organic and metal toxicants, e.g., it is very sensitive to copper. Bioassay studies in 1982 indicated that this estuary is not generally suitable for the toxic dinoflagellate Gonyaulax tamarensis (=Alexandrium tamarense) (Mahoney et al., 1988).

Comparison of A. anophagefferens incidence from a non-bloom year and a bloom year suggests that A. anophagefferens regulation can be general from Barnegat Bay south to Cape May at least, and not specific to the Barnegat Bay-Little Egg Harbor-Great Bay system where it flourishes best. The western Long Island incidence data are limited but serve to begin to close an information gap. A possible explanation for the higher June, 2001 levels in western Long Island coastal ocean sites (74. 75, 76) relative to estuarine sites is that a minor bloom (cell concentration to 4.6 x 105 cells ml-1; Nuzzi, unpublished data) occurred at the same time in mid- to western Great South Bay and bloom water may have flowed westward from Fire Island Inlet. Levels of A. anophagefferens in Long Island and New Jersey coastal ocean waters were roughly comparable.

Recommendations

Because an intense bloom of A. anophagefferens is not necessary for detrimental effects, in addition to bloom centers monitoring should be conducted in areas that have history or apparent potential for supporting levels of the species > 3.5 x 104 < 2.0 x 105 cells ml-1.

The Hudson-Raritan estuary and western Long Island bays apparently have never experienced brown tides but are proximal to areas where brown tides recur. Because at least some of these waters characteristically support high primary productivity and factors regulating bloom occurrence may change, regular survey of these waters is recommended .

IN MEMORIAM

One of the authors, Christine Zetlin, is recently deceased. She is missed by her colleagues. Typical of her research dedication, she contributed to this report while seriously ill.


ACKNOWLEDGMENTS

We thank the personnel of the Bureau of Marine Resources, Suffolk County, NY, Department of Health Services, particularly R. Nuzzi and J. Bredemeyer, for training James J. Howard Marine Sciences Laboratory personnel in A. anophagefferens enumeration and contributing some of the incidence data. We thank members of the USEPA, Region II, Surveillance and Monitoring Branch, for helicopter water sample collections over the course of the study. E. Cosgrove, Monmouth County, NJ, Department of Health, provided some of the water temperature data. We thank K. Wendling for technical assistance and A. Kalbach for assistance with figure preparation.


LITERATURE CITED

ANDERSON, D.M., D.M. KULIS, and E.M. COSPER. 1989. Immunofluorescent detection of the brown tide organism Aureococcus anophagefferens. Pp. 213-228 in Cosper, E.M., V.M. Bricelj, and E.J. Carpenter (eds). Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms. Springer-Verlag, Berlin.

ANDERSON, D. M., B. A. KEAFER, D. M. KULIS, R. M. WATERS AND R. NUZZI. 1993. An immunofluorescent survey of the brown tide chrysophyte Aureococcus anophagefferens along the northeast coast of the United States. J. Plank. Res. 15:563-580.

BRICELJ, V.M. AND S.H. KUENSTER. 1989. Effects of the "brown tide" on the feeding, physiology and growth of bay scallops and mussels. Pp. 491-508 in Cosper, E.M., V.M. Bricelj, and E.J. Carpenter (eds.). Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms. Springer-Verlag, Berlin.

BRICELJ, V. M. AND D. J. LONSDALE. 1997. Aureococcus anophagefferens: Causes and consequences of brown tides in U. S. mid-Atlantic coastal waters. Limnol. Oceanogr. 42 (5, part 2): 1023-1038.

BRICELJ, V. M., S. P. MACQUARRIE AND R. A. SCHAFFNER. 2001. Differential effects of Aureococcus anophagefferens isolates ("brown tide") in unialgal and mixed suspensions on bivalve feeding. Mar. Biol. 139:605-615.

DZURICA, S., C. LEE, E. M. COSPER, AND E. J. CARPENTER. 1989. Role of environmental variables, specifically organic compounds and micronutrients, in the growth of the chrysophyte Aureococcus anophagefferens. Pp. 229-252 in Cosper, E. M., V. M. Bricelj, and E. J. Carpenter (eds.). Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms. Springer-Verlag, Berlin.

MAHONEY, J. B., D. HOLLOMON, AND R. WALDHAUER. 1988. Is the lower Hudson-Raritan estuary a suitable habitat for Gonyaulax tamarensis? Mar. Ecol. Prog. Ser. 49:179-186.

MAHONEY, J. B., D. JEFFRESS, J. BREDEMEYER, P. OLSEN. 1999. Aureococcus anophagefferens brown tide in New Jersey. Pp. 76-81 in Martin, J. L., and K. Haya (eds.) Proceedings of the Sixth Canadian Workshop on Harmful Marine Algae. Canadian Technical Report of Fisheries and Aquatic Sciences No. 2261.

NUZZI, R. 1995. The brown tide - an overview . Pp. 13-23 in McElroy, A. (ed.). Proceedings of The Brown Tide Summit. Publ. No. NYSGI-W-95-001, New York Sea Grant Institute.

NUZZI, R., P. OLSEN, J. B. MAHONEY, AND G. ZODL. 1996. The first Aureococcus anophagefferens brown tide in New Jersey. Harmful Algae news, IOC Newsletter No. 15.

OLSEN, P. 1989. Development and distribution of a brown water algal bloom in Barnegat Bay, New Jersey, with perspective on resources and other red tides in the region. Pp. 189-212 in Cosper, E. M., V. M. Bricelj, and E. J. Carpenter (eds.) Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms. Springer-Verlag, Berlin.

OLSEN, P. S. AND J. B. MAHONEY. 2001. Phytoplankton in the Barnegat Bay-Little Egg Harbor estuarine system: species composition and picoplankton bloom development. J. Coast. Res. Special Issue No. 32: 115-143.

O'REILLY, J., J. P. THOMAS AND C. EVANS. 1976. Annual primary production (nanoplankton, netplankton, dissolved organic matter) in the Lower New York Bay. Pp. 1-39 in McKeon, W. H, and G. J. Lauer (eds.) Proceedings of the Fourth Symposium on Hudson River Ecology. Hudson River Environmental Society, Inc., New York.

SIEBURTH, J. McN., P.W. JOHNSON, AND P.E. HARGRAVES. 1988. Ultrastructure and ecology of Aureococcus anophagefferens gen. et sp. nov. (Chrysophyceae): the dominant picoplanker during a bloom in Narragansett Bay, Rhode Island, Summer 1985. J. Phycol. 24:426-425.

SMAYDA, T. J. AND T. A. VILLAREAL. 1989. The 1985 "Brown Tide" and the open phytoplankton niche in Narragansett Bay during summer. Pp. 159-188 in Cosper, E. M., V. M. Bricelj, and E. J. Carpenter (eds.) Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms. Springer-Verlag, Berlin.

STEELE, R. L., L. C. WRIGHT, G. A. TRACEY AND G. B. THURSBY. 1989. Brown tide bioassay: growth of Aureococcus anophagefferens Hargraves et Sieburth in various known toxicants. Pp. 253-264 in Cosper, E. M., V. M. Bricelj, and E. J. Carpenter (eds.) Novel Phytoplankton Blooms: Causes and Impacts of Recurrent Brown Tides and Other Unusual Blooms. Springer-Verlag, Berlin.

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