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U.S. Geological Survey Open-File Report 00-306: Chapter 8

Holocene Dinoflagellate Cysts from Marion-Dufresne Core MD99-2209, Chesapeake Bay

by Stacey Verardo
Atlantic GeoServices Inc., Oakton, Virginia 22124
c/o United States Geological Survey, Reston, Virginia 20192


Dinoflagellate cysts from Marion-Dufresne core MD99-2209 in Chesapeake Bay were analyzed to examine spatial and temporal trends in dinoflagellates during the Holocene. Results show that during the Early Holocene about 6600-5900 yr BP (1700-900 cm) there are centennial-scale oscillations in the mean abundance of several species of Spiniferites, (i.e. S. ramosus, S. bulloideus, and S. mirabilis), Operculodinium centrocarpum, Lingulodinium macherophorum and Polysphaeridium zoharyi. These shifts probably signify variations in salinity and water temperature. During the Late Holocene ~ 2300-present yr. BP (800-0 cm) there was a progressive rise in relative abundance of Spiniferites spp. coincident with a decrease in relative abundance in most other species. One of the most important changes in Chesapeake Bay dinoflagellates over the past two millennia has been the increase in abundance of S. mirabilis during the past two centuries, an event which signifies the impact of anthropogenic activities on Bay phytoplankton in the watershed.


Dinoflagellates are organic-walled single-celled organisms that produce cysts within motile dinoflagellate theca (Fensome and others, 1996) and occur in marine, brackish, and freshwater environments. Living dinoflagellates may be autotrophs, heterotrophs or symbionts. Photosynthetic forms (autotrophs) account for approximately 50% of the dinoflagellate genera. Free living genera are primary producers in the oceans thereby comprising an important component of marine phytoplankton.

Fossil remains of dinoflagellates consist primarily of the nonmotile or cyst stage. Dinoflagellate cysts are common in most sediment samples from the Chesapeake Bay. They are useful in documenting temporal trends in Chesapeake phytoplankton during climatic changes in the Holocene, as well as during the postcolonial period of 19th century agricultural land clearance and 20th century fertilizer application and urbanization.


Core MD99-2209 was recovered from the Chesapeake Bay off the Rhode River and split into archive and working core sections. The working sections were lithologically described and sampled on the ship for dinoflagellate cysts at 10 cm intervals from the surface to a depth of 1700 cm. The samples discussed in this study are the same samples used for palynological analysis discussed by Willard and Korejwo (Chapter 7, this volume).

Dried sediment samples, weighing between 4.0 and 7.0 grams, were prepared for palynological analysis by decalcifying with dilute (i.e. 10%) hydrochloric acid and then neutralizing (to a pH of 7) with deionized water to remove carbonates. After neutralization, hydrofluoric acid (52% by volume) was added to each sample and the resulting mixture was allowed to rest at room temperature for ~48 hours. After resting, the samples were rinsed with deionized water into 50 milliliter (ml) centrifuge tubes. A 10 ml solution of acetic anhydride and sulfuric acid, in a ratio of 9:1, was added to each sample. Samples were heated in a water bath at 100°C for 10 minutes after which, 25 ml glacial acetic acid was added to each sample to arrest the reaction. The samples were then treated with 25 ml solution of 10% potassium hydroxide and heated to 70°C in a water bath.

The samples were then centrifuged with a Damer/IEC Model K centrifuge at 1500 rpm and washed until neutralized to a pH of 7. If the samples contained sand, heavy minerals, or organic material, they were centrifuged at 500 rpm for 20 minutes and then at 1500 rpm for 10 minutes with 3 ml zinc chloride, a heavy liquid with a specific gravity of 2.1. After neutralization, the samples were sieved through 1249 micron (µm) and 10µm nylon mesh sieves to remove plant material and clay-sized particles. The residue was mixed with warm (i.e. 5°C) glycerin jelly and placed on microscope slides for final preparation.

Two drops of the residue and glycerin jelly mixture was dropped onto a clean slide and placed on a warming table at ~40°C. The coverslips were carefully placed on the slides and allowed to cure for 48 hours on the warming table. The slides were cured upside-down (by supporting the edges of the slides with thin wood dowels) to bring the fossil cysts closer to the coverslip, making it easier to focus. The coverslip was then outlined with clear acrylic to seal the slide and each slide was labeled with location, sample date, and depth.

The slides were mounted on a binocular microscope to identify and count a minimum of 300 dinoflagellate cysts for meaningful statistical analysis (Imbrie and others, 1973). Scanning Electron Micrographs were taken of individual dinoflagellates to insure taxonomic identity (Verardo, 1999). The relative abundance (i.e. percent of total number of individuals) of each species was calculated by dividing the number of individuals by the total number of individuals in each sample.

The age model for MD99-22099 was developed from radiocarbon dating (see Colman and others, Chapter 6, this volume) and pollen biostratigraphy (Willard and Korejwo, Chapter 7, this volume). One datum of particular importance when interpreting dinoflagellate trends is the agricultural horizon, which occurs at ~250-260 cm in core MD99-2209. This horizon refers to the sharp rise in Ambrosia (ragweed) pollen to greater than 10% of the total pollen assemblage that occurred during the 1800's (Brush and others, 1982). Willard and others (in prep.) discuss recent data on the timing and impact of 19th century land clearance on Chesapeake Bay and its watershed.


Figure 8.1 plots the relative frequencies (% abundance) of six dinoflagellate taxa in Chesapeake Bay core MD99-2209. The dominant dinoflagellate genus in the Chesapeake Bay is Spiniferites spp., which includes in this region representatives of the subtropical species S. ramosus and S. bulloideus (S. mirabilis was counted separately, see below). Today, these species of Spiniferites occur predominantly in oceanic environments along the southeastern United States, as well parts of the Caribbean (Wall and others, 1977, Harland, 1983; de Vernal and others, 1992).

The mean abundance of Spiniferites cysts in MD99-2209 varies from <30 to ~60% between the bottom of the core (~1700 cm) and the unconformity near 800 cm. Above the unconformity the abundance of Spiniferites progressively increases, reaching a peak above 80% near 200 cm, above which it varies between 50 and 80%. These variations in relative abundance presumably signify changes in temperature and salinity, although we cannot yet quantify these factors for this site. It is noteworthy that the relatively high proportions of Spiniferites in the uppermost 200 cm, which corresponds to the past 150 years, is reminiscent of the dominance of this genus between 1600 and 1720 cm.

One species in particular, Spiniferites mirabilis, seems to be an excellent indicator of the negative impact of 19th century agriculture and 20th century fertilizer application had on Bay dinoflagellates. Prior to the 19th century, this species varied from <4-7% of the total dinocyst assemblage, with the exception of an interval between 600-700 cm (about 1300-1600 yr BP). Pre-historical low abundance of S. mirabilis contrasts sharply with its abundance since the middle 1800's, when it increased to 10-15%, occasionally reaching 20% during the 1900's. Harland (1983) and Turon (1984) present evidence that S. mirabilis can tolerate increased nutrient concentrations of phosphate and nitrate and warm surface waters. Recent studies of the modern distribution of S. mirabilis in coastal bays and estuaries along the Atlantic and Gulf coasts of North America support the idea that this species is extremely tolerant of high nutrient levels, reduced and variable salinity, high turbidity, and perhaps reduced levels of dissolved oxygen (Verardo, unpublished data; Willard and others, in prep.). The results from MD99-2209 demonstrate that during the last 30 years, S. mirabilis has reached unprecedented levels of abundance in Chesapeake Bay, probably related to changing environmental conditions.

The second most abundant dinoflagellate genus in MD99-2209 is Operculodinium centrocarpum. O. centrocarpum has a modern distribution associated with estuarine and coastal waters in mild to cool temperate regions (Wall and others, 1977; Harland, 1983; de Vernal and others, 1992; Brenner, 1998). This species comprised 5 to 30% of early Holocene assemblages between 1600 and 1400 cm. The abundance then decreased to 10% before increasing to 25% by 900 cm. O. centrocarpum maintained a steady relative abundance of 5-15% for much of the past 2000 years (above 800 cm) although a small net decline can be observed and increased variability above the 200 cm level. Other species, such as Polysphaeridium zoharyi and Lingulodinium macherophorum are presently both found along the southeastern seaboard of the United States and the Caribbean and are considered tropical-subtropical marine species (Wall and others, 1977; Harland, 1983; Edwards and Anderle, 1992). Both of these species exhibit oscillating abundances between 5-10% during the early Holocene (between 1700 cm and 900 cm).

One of the noteworthy results to emerge from this preliminary data is the progressive decrease in the relative abundances of Polysphaeridium zoharyi, Lingulodinium macherophorum and Nematophaeropsis spp. in the last ~2300 years of Late Holocene deposition (the interval 800 cm to 0 cm). Nematophaeropsis spp., a genus found in the North Atlantic associated with the cold North Atlantic Gyre (Wall and others, 1977; Harland, 1983; Edwards and Anderle, 1992), decreased to ~1% at the surface. The only taxonomic groups that did not decrease in abundance during the Late Holocene were Spiniferites spp. and S. mirabilis, which presently is the dominant taxon in the Bay. These trends are also observed in dinoflagellate assemblages from other Chesapeake Bay cores (Verardo and others, 1998; Willard and others, in prep.) and suggest a diminished phytoplankton biodiversity during the post-Colonial period.


The trends in dinoflagellate cysts in Chesapeake Bay appear to be consistent with other evidence indicating both climatic and anthropogenic influences on the Bay's environments. However, because dinoflagellate cysts are resistant to degradation in sediments and might also be reworked from Cenozoic deposits that outcrop around Chesapeake Bay, it is important to be certain those cysts found in Holocene sediment cores are in situ and not transported from older sediments.

There are several reasons to support the argument that the cysts recovered from MD99-2209 and other Holocene sediments from Chesapeake Bay cores represent autochthonous assemblages that inhabited Chesapeake Bay. First, the excellent preservation of cysts (Verardo, 1999), including coloration similar to cysts from modern sediments, suggests they have not been transported. Second, although species such as P. zoharyi occur in sediments near the Bay (Edwards, 1986), P. zoharyi is also a component of the modern assemblage. Extinct Miocene species have rarely been encountered in the several hundred Holocene dinocyst samples from the Chesapeake Bay. Third, other microfaunal and floral paleoecological indicators (Cooper and Brush, 1991; Cronin and others, 2000) show concordant trends with those of the dinoflagellates, suggesting the entire ecosystem, including phytoplankton, respond to environmental change. Fourth, P. zoharyi and other Chesapeake Bay dinoflagellate taxa typically live in marine environments, phytoplanton populations could easily enter the Bay at its mouth where deep saline water flows into the main channel from the continental shelf. In sum, there is no evidence for significant reworking of Cenozoic dinoflagellates in the Marion-Dufresne and other Chesapeake Bay sediment cores.


Dinoflagellate cyst assemblages from MD99-2209 reveal both natural, pre-colonial changes in Chesapeake Bay phytoplankton and post-colonial influence of land use changes. The increase in the mean abundance in Spiniferites spp. over the past 2300 years suggests changes in salinity and perhaps temperature. The rise in S. mirabilis from background levels of <5% to 10-20% implies a net decrease in dissolved oxygen and increase turbidity in the Chesapeake Bay during the past two centuries. These unprecedented changes in the dinoflagellate cyst assemblages are clearly linked to land clearing, agriculture, and other anthropogenic activities in the watershed, which led to increases in sediment and nutrient loads. However, there are also frequent large oscillations in the percent abundance of several dinoflagellate taxa during both the early and the late Holocene. These fluctuations in Bay phytoplankton imply dynamic short-term variability in bay environments that must be related to natural variations in hydrological conditions. Regional climatic variability could, in theory, influence Bay salinity through its impact on precipitation and tributary discharge and water temperatures. Therefore, both human-induced and natural factors areresponsible for observed changes in dinoflagellate cyst diversity and assemblage composition in Chesapeake Bay.


I thank Lucy Edwards for her help with dinoflagellate taxonomic. I am also grateful to Lisa Weimer, Tom Sheehan and James Murray for producing quality slides as well as Robert Wagner and Moira Slattery for processing samples.


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