Ceratium in Fire Island Inlet, Long Island, New York (1971-1977)

Ceratium in Fire Island Inlet, Long Island, New York (1971-1977)Author(s): Sylvia S. WeaverSource: Limnology and Oceanography, Vol. 24, No. 3 (May, 1979), pp. 553-558Published by: American Society of Limnology and OceanographyStable URL: http://www.jstor.org/stable/2835616 .

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Notes 553

SIOLI, H. 1964. General features of the limnology of Amazonia. Int. Ver. Theor. Angew. Limnol. Verh. 15: 1053-1058.

. 1968. Principal biotypes of primary production in the waters of Amazonia. Proc. Symp. Rec. Adv. Trop. Ecol. 2: 591-600.

SMITH, G. M. 1933. Freshwater algae of the United States. McGraw-Hill.

STRICKLAND, J. D., AND T. R. PARSONS. 1972. A practical handbook of seawater analysis, 2nd ed. Bull. Fish Res. Bd. Can. 167.

THOMASSON, K. 1971. Amazonian algae. Inst. R. Soc. Nat. Belg. Mem. 2: 1-57.

VOLLENWEIDER, R. A. 1974. A manual of methods for measuring primary production in aquatic environments, 2nd ed. IBP Handbook 12. Blackwell.

WETZEL, R. G. 1975. Limnology. Saunders. WHITFORD, L. A., AND G. J. SCHUMACHER. 1973.

A manual of freshwater algae. Sparks. WILLIAMS, P. M. 1968. Organic and inorganic con-

stituents of the Amazon River. Nature 218: 937-938.

Submitted: 30 March 1978 Accepted: 12 October 1978

Limnol. Oceanogr., 24(3), 1979, 553-558 ?) 1979, by the American Society of Limnology and Oceanography, Inc.

Ceratium in Fire Island Inlet, Long Island, New York (1971-1977)

Abstract-Documentation of the occurrence of the dinoflagellate Ceratium tripos (0. F. Muller) Nitzsch, at Fire Island Inlet from 1971 through 1977 provides a perspective for its 1976 mid-Atlantic coastal bloom.

The dinoflagellate Ceratium tripos occurred in very large numbers off the entire U.S. mid-Atlantic coast from Febru- ary through July 1976. Bulloch (1976) and Malone (unpubl.) suggested that this bloom contributed to the anoxia and sub- sequent fish kill off the New Jersey coast during this period. My observations since 1971 in the nearshore waters of Fire Is- land Inlet, off the south shore of Long Island (Fig. 1), show that while the mag- nitude of the 1976 bloom was extraordi- nary, C. tripos has occurred each year with peaks in April and May. Here I re- port its occurrence over the last 7 years.

Zooplankton and phytoplankton samples were collected biweekly at Oak Beach in Fire Island Inlet. The techniques are described by Weaver and Hirshfield (1976), who studied bay and ocean stations in the area from 1971- 1974. They found by comparing percent frequency of occurrence, relative abundance, rank-order correlation, and indi- cator species that two plankton popula-

tions (estuarine-bay and nearshore-ocean) could be monitored at one station (Oak Beach) by sampling at midtide after slack current on both ebb- and floodtides. Here I deal with the nearshore-ocean samples only, since C. tripos was not found in bay-estuarine samples.

Bottle and net (160-Am mesh) surface samples (<1 m) were taken simulta- neously, the net set up off the dock to take advantage of the current for a 10-min stationary "tow" (Fish 1925). Current speed, measured by tethered drogue (Horrer 1967), averaged 0.5 m s-1. Tidal range averaged 1.3 m. Depth at the sta-

4100-

40A 30- Inlet

N E JE EY

740?00 73 00'

Fig. 1. New York Bight, showing study area.



554 Notes

25-__ _ _

20 TEMPER TURE o15-

0 0_

5-

32 SALINI Y 0 31 -'fV ~ " \

500-

CERATI M TRIPO~

100-

50_

4+2 5-

0.5-

0.1- Zt III ln 1 |1 monthFA M J F M A M J M J J F MA M J J A M

1 973 1974 1975 1976 1977 Fig. 2. Ceratium tripos, temperature, and salinity at Fire Island Inlet (1973-1977).



Notes 555

tion varied from 3 to 4 m annually due to shifting sands. The water column was mixed. Salinity was calculated from temperature and density measurements (Mangelsdorf 1967).

Live observations were made of both net and bottle samples (centrifuged). Pre- served phytoplankton (Lugol's merthio- late: Weber 1968) was counted in a Palm- er and Maloney (1954) cell at 400x from bottle samples (300 ml) after progressive settling. This tally excluded C. tripos and Coscinodiscus centralis, which were counted in a Sedgwick-Rafter cell at lOOx with the Formalin-preserved net plankton because of their large size (200- 300 Am). These organisms may escape routine notice because they are too large to be counted as numerical dominants with the usual small-scale phytoplankton bottle sampling and counting techniques. Conversely, they may be ignored in zooplankton net hauls because they are, tax- onomically speaking, phytoplankters. Al- though relative percentages can be derived accurately, net tows are unreli- able in terms of absolute numbers. Therefore, I compared counts of Cerati- um for all 1976 net samples (average of three Sedgwick-Rafter counts) and for all 1976 bottle grabs (average of three Palm- er-Maloney whole cell counts). Results confirmed a net factor of 100, by which absolute cell counts of Ceratium liter-' could be reliably extrapolated from net samples collected before 1976 and therefore tallied only in the Sedgwick-Rafter cell. This large net factor was corrobo- rated in the analysis of replicate samples examined at the height of the bloom (17 May 1976) to test for patchiness (unob- served) and is probably due to rapid net- clogging by this species.

Ceratium tripos was present in the nearshore-ocean community during late winter and spring in each year of the quantitative study (1973-1977) at Fire Is- land Inlet (Fig. 2). Maximum abundances varied considerably from a low of 95. liter-' in 1977 to a high of 42,200 in 1976. Qualitative observations also

showed peaks in February and April 1972 and in March and May 1971.

Water temperature may be an indirect determinant of Ceratium species varia- tion, abundance, or both. Ceratium is found at Fire Island Inlet within the range of 2?-21?C; salinity (29-33%o) does not appear to be limiting (Fig. 2). Graham (1941), Nordli (1953), Hulburt and Rod- man (1963), Mulford (1963), and others have also found C. tripos within a wide range of temperatures and salinities. But the onset and termination of C. tripos blooms do not appear to be influenced by temperature.

The 1973 and 1975 winter temperatures before the appearance of Ceratium were higher than in 1974 and 1976. Ce- ratium did not appear until the water temperature reached 12?C in 1973 and 18?C in 1975. The dominant species in these years were Ceratium longipes, Ce- ratium gracile, and Ceratium massiliense, as well as C. tripos. It seems like- ly that the species variety present in those years indicated some different con- dition or body of water, expressed as a temperature response. Bigelow (1924) described C. longipes as a "warm water" species in the Gulf of Maine; C. gracile and C. massiliense are "tropical" species (Graham 1941; Mulford 1963). Fornshell et al. (unpubl.) found Cape Hatteras to be a dividing point for Ceratium species in 1976: C. tripos and Ceratium macroce- ros to the north, C. massiliense and Ce- ratium trichoceros to the south.

The temperature adaptability of C. tripos is revealed in 1974 and 1976. Early winter temperatures were lower than in 1973 and 1975, with a seasonal low of 20C in March 1974 and at the end of January in 1976. The maximum Ceratium count occurred in May 1974 at 140C and in May 1976 at 150C. In every year all Ceratium species declined soon after the temperature reached 180C.

Unusually low water temperatures (to -20C) in January and February may have delayed the appearance of C. tripos in 1977 or may have been more favorable to



556 Notes

Table 1. Phytoplankton dominants associated with Ceratium blooms at Fire Island Inlet (1973-1977).

Cells liter1

Date Species % No. 103

26 May 73 Chaetoceros perpusillus 21 3 Asterionella japonica 15 2

12 Jun 73 Thalassiosira nana 36 25 Ceratium lineatum 26 18

26 Mar 74 Thalassiosira nordenskioldii 35 375 Plagiogramma brockmanni 21 225

13 Apr 74 A. japonica 46 455 P. brockmanni 16 158

28 Apr 74 P. brockmanni 39 105 11 May 74 Cerataulina bergoni 76 205 26 May 74 T. nana 38 114

P. brockmanni 22 66 24 May 75 Prorocentrum minimum 34 37

T. nana 24 26 14 Jun 75 Leptocylindrus danicus 58 162

Rhizosolenia delicatula 15 42 15 Feb 76 Skeletonema costatum 75 2,775 2 Mar 76 S. costatum 51 1,275

Nitzschia seriata 39 975 15 Mar 76 N. seriata 83 1,245 1 Apr 76 N. seriata 36 504

Peridinium triqueta 24 336 15 Apr 76 P. triqueta 50 650

N. seriata 38 494 1 May 76 S. costatum 28 84

N. seriata 22 66 17 May 76 R. delicatula 47 376

C. perpusillus 35 280 22 May 76 S. costatum 49 294 5 Jun 76 T. nana 59 236

24 Jun 76 no dominants 3 Jul 76 S. costatum 64 192

23 Apr 77 P. minimum 29 55

the extended bloom of the haptophycean Phaeocystis pouchetti (3,515 colonies* liter-', 500 cells colony-' on 16 April), which may in turn inhibit Ceratium. This pattern was repeated in 1978. Ex- cept in 1974, Ceratium and Phaeocystis have never been found together in Fire Island Inlet; their possible interaction has not been investigated.

I saw no consistent phytoplankton species association (Table 1) except that Ceratium follows C. centralis and Phaeocystis seasonally and that early 1974 and 1976 were characterized by ex- ceptionally heavy winter blooms of Skel- etonema costatum, followed in March 1974 by Thalassiosira nordenskioldii and in March 1976 by Nitzschia seriata. Yet 1977 also produced a heavy winter

bloom of S. costatum, while all Ceratium species were sparse. In addition, the an- nual bloom of Phaeocystis in 1977 lasted much longer than usual, accompanied by abundant Leptocylindrus minimus. In 1973 and 1975, S. costatum was scarce in the winter plankton and the C. tripos count was comparatively low (Fig. 2).

No zooplankton predation or endopar- asitism was observed. Calanoid copepods (primarily Acartia clausii), Oithona sim- ilis, and spionid larvae were the most abundant zooplankton from January through June. Hydromedusae (Sarsia, Margelopsis, Obelia) swarm in April. In 1973 and 1975, the cladoceran Evadne nordmanni was a dominant in May.

Although direct dependence of Cera- tium on temperature cannot be dem-



Notes 557

onstrated, water temperature as it may reflect large-scale wind and water movements could account for Ceratium species variety and for the seasonal onset and duration of C. tripos at Fire Island Inlet and all along the east coast in 1976. Prevailing winds in 1976 (U.S. Dep. Commerce 1976: Local climatological data) shifted earlier than usual from north to south, leading to a more westerly flow overall. When we consider the coastwide occurrence of Ceratium, this could have caused a concentration of C. tripos par- allel to the south shore of Long Island (cf. Rounsefell and Dragovich 1966), ac- counting for unusually high numbers in Fire Island Inlet in March and April compared with other years. Increases in cell density paralleled those observed in the apex of the New York Bight (Malone 1977) and on the shelf south of Long Island (Walsh et al. 1978). At Fire Is- land Inlet, numbers increased from <100 liter-1 in January to 22,000 liter-t at the end of March, remained more or less stable through April and May, reached a maximum of 43,000 liter-t at the end of May, and declined again to <100 liter-1 by mid-July (Fig. 2). The bulk of the Ceratium bloom in the New York Bight after stratification in April and May was found at or below the thermocline (Malone unpubl.), but local storms off the south shore of Long Island kept the water column mixed through May (Walsh et al. 1978). This was reflected in the peak of the Ceratium bloom in Fire Island Inlet at the end of May. In June, the Ceratium population on the shelf south of Long Island aggregated below the thermocline at the 1-10% light layer (Malone unpubl.). Prevailing south- southwesterly winds in June and July and the consequent movement of bottom water inshore probably accounted for the continued abundance of C. tripos in the well mixed waters of Fire Island Inlet through June and, briefly, in July. Walsh et al. (1978) pointed out that such "wind events" may ultimately control all phytoplankton dynamics along the entire mid-Atlantic shelf.

It is perhaps indicative of a twofold pulse of Ceratium that morphological variations (longer or shorter apical and antapical horns) were far more abundant in the C. tripos population through mid- May, probably indicating different growth stages. From mid-May on, the morphol- ogy of individuals was uniform, suggest- ing a nongrowing population-one that might have originated from an aggrega- tion below the compensation level.

Thus, while the occurrence of C. tripos in 1976 shelf waters was not unusual (Mandelli et al. 1970; Smayda 1973; Weaver and Hirshfield 1976; Marshall 1976), its numbers were. This may be explained by the fortuitous combination of winds, water temperature, and oppor- tunistic species physiology enabling Ce- ratium to survive for long periods at low light levels. Further field observations and experiments should clarify this.

Insofar as Ceratium dominance in late winter-early spring was observed at Fire Island in 1976, and in the perspective of observations there for the last 7 years, it seems that monitoring of nearshore stations is a useful adjunct to ocean-going research. Such stations can be active year round and, strategically located, could provide an "early warning" network as well as continuing background data for just such events as the bloom of C. tripos in the New York Bight in 1976.

Sylvia S. Weaver 50 West 72nd St. New York, New York 10023

References BIGELOW, H. B. 1924. Plankton of the offshore

waters of the Gulf of Maine. Bull. Bur. Fish. Wash. 40: 407-417.

BULLOCH, D. K. 1976. Ocean kill in the New York Bight-Summer 1976. Underwater Nat. 10(1): 4-12.

FISH, C. J. 1925. Seasonal distribution of the plankton of the Woods Hole Region. Bull. Bur. Fish. Wash. 41: 91-179.

GRAHAM, H. W. 1941. An oceanographic consid- eration of the dinoflagellate genus, Ceratium. Ecol. Monogr. 11: 99-116.

HORRER, P. L. 1967. Methods and devices for measuring currents, p. 80-89. In G. H. Lauff [ed.], Estuaries. Publ. Am. Assoc. Adv. Sci. 83.



558 Notes

HULBURT, E. M., AND J. RODMAN. 1963. Distri- bution of phytoplankton species with respect to salinity between the coast of southern New England and Bermuda. Limnol. Oceanogr. 8: 263-267.

MALONE, T. 1977. Light-saturated photosynthesis by phytoplankton size fractions in the New York Bight, USA. Mar. Biol. 42: 281-292.

MANDELLI, E. F., P. R. BURKHOLDER, T. E. DOH- ENY, AND R. BRODY. 1970. Studies of primary productivity in coastal waters of southern Long Island, New York. Mar. Biol. 7: 153-160.

MANGELSDORF, P. C., JR. 1967. Salinity measurements in estuaries, p. 71-79. In G. H. Lauff [ed.], Estuaries. Publ. Am. Assoc. Adv. Sci. 83.

MARSHALL, H. G. 1976. Phytoplankton distribution along the eastern coast of the USA. 1. Phy- toplankton composition. Mar. Biol. 38: 81-89.

MULFORD, R. A. 1963. Distribution of the dinoflagellate genus Ceratium in the tidal and offshore waters of Virginia. Chesapeake Sci. 4: 84-89.

NORDLI, E. 1953. Salinity and temperature as con- trolling factors for distribution and mass occurrence of Ceratia. Blyttia 1 1: 16-18.

PALMER, C. M., AND T. E. MALONEY. 1954. A new counting slide for nannoplankton. Am. Soc. Limnol. Oceanogr. Spec. Publ. 21. 6 p.

ROUNSEFELL, G. A., AND A. DRAGOVICH. 1966. Red tide and oceanographic factors. Bull. Mar. Sci. 16: 404-422.

SMAYDA, T. J. 1973. A survey of phytoplankton dynamics in the coastal waters from Cape Hatter- as to Nantucket, p. 1-99. In Coastal and offshore environmental inventory. Cape Hatteras to Nantucket Shoals. Univ. R.I. Mar. Publ. Ser. 2.

WALSH, J. J., AND OTHERS. 1978. Wind events and food chain dynamics within the New York Bight. Limnol. Oceanogr. 23: 659-683.

WEAVER, S. S., AND H. I. HIRSHFIELD. 1976. The delineation of two plankton communities from one sampling site (Fire Island Inlet, Long Is- land, N.Y.). Mar. Biol. 34: 273-283.

WEBER, C. I. 1968. The preservation of phytoplankton grab samples. Trans. Am. Microsc. Soc. 87: 70-81.

Submitted: 22 November 1977 Accepted: 24 July 1978

Limnol. Oceanogr., 24(3), 1979, 558-562 ? 1979, by the American Society of Limnology and Oceanography, Inc.

The toxic effect of copper on Oscillatoria (Trichodesmium) theibautiil Abstract-[14C]CO2 fixation in Oscillatoria

theibautii decreased in response to copper at low cupric ion activities (10-10 M) in test media containing Tris as a copper chelator and to low concentrations (10-8 M) of total added copper in filtered seawater. This sensitivity could be important to natural systems as well as laboratory and shipboard cultures.

Oscillatoria (Trichodesmium) theibautii is a marine blue-green alga common in the Sargasso and Caribbean seas. It can fix elemental nitrogen and could provide an input of this nutrient into those waters (Goering et al. 1966; Carpenter and McCarthy 1975). Study of 0. theibautii has been restricted because it is very difficult to keep viable for any sig- nificant period (Moreth 1970). Culturing

1 Research was funded by NSF grant DES 75- 15023 to F. M. M. Morel and ship time was provid- ed through NSF grant OCE 75-14781 to J. J. Mc- Carthy.

attempts have had only limited success, of up to 3 months (Carpenter and Mc- Carthy 1975; J. Waterbury pers. comm.).

The important parameter of copper tox- icity to phytoplankton is the cupric ion activity (Sunda and Guillard 1976). Dif- ferent species show different sensitivi- ties when grown in the artificial media "Aquil" (Morel et al. 1975) with the mid- point of the toxic response at 10-10.5 M cupric ion activity for Gonyaulax tamar- ensis (Anderson and Morel 1978) and at 10-8 M cupric ion activity for Skeletone- ma costatum (SKEL) (Rueter 1977; Mo- rel et al. 1978). Some freshwater blue- green algae are also reported to be very sensitive to copper (Horne and Goldman 1974). Species with well characterized responses to cupric ion activity have been used as bioassay organisms to de- termine the copper complexation capac- ity of natural seawater (Davey et al. 1973).



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Ceratium in Fire Island Inlet, Long Island, New York (1971-1977)