Tag: salt marsh ecology

The Hidden City in the Grass

How seagrasses and marsh grasses—and the animals within them—build the marshes of Onslow County

In Onslow County’s estuarine marshes, the best time to understand how the landscape works is when the water pulls back. As tides drain from creeks and shallow flats, patterns begin to emerge—where water lingers, where it moves easily, and where it hesitates. These patterns are not random. They reflect the combined influence of plants, animals, and sediments continually reshaping the boundary between land and sea.

Like the microscopic shells of foraminifera preserved in sediment, marsh and seagrass communities record environmental conditions. But unlike the past locked in mud, these systems are alive, constantly negotiated by plants, grazers, predators, and microbes.

From permanently submerged seagrass beds to the highest marsh edge, each elevation zone in Onslow County is maintained not just by vegetation, but by species that actively regulate growth, chemistry, and water flow.

Subtidal shallows: seagrass beds maintained by grazers

In the shallow, light-penetrated waters of the New River Estuary and protected soundside areas, seagrass beds form underwater meadows that stabilize sediments and provide nursery habitat for fish and invertebrates. Species present or expected in Onslow County waters include eelgrass (Zostera marina), shoalgrass (Halodule wrightii), and widgeongrass (Ruppia maritima) (Mallin, 2000; Orth, 1984).

Seagrass blades rapidly accumulate epiphytic algae and microbial films. Without constant grazing, this layer can block light and suppress photosynthesis. Amphipods, isopods, and small gastropods act as continuous maintenance crews, grazing epiphytes and preventing them from overwhelming the plants themselves (Orth & van Montfrans, 1984; Valentine & Duffy, 2006).

Experimental studies show that when these grazers are removed, seagrass condition declines even under favorable light conditions, demonstrating that plant survival depends as much on animal activity as on physical environment (Duffy et al., 2015). Beneath the canopy, burrowing worms and bivalves recycle nutrients and oxygenate sediments, preventing organic matter from accumulating around roots (Orth, 1984).

In this zone, seagrass persists because grazers keep blades clean and sediments breathable—a cooperative system built on constant biological upkeep.

Gammarus mucronatus, a common amphipod grazer on eelgrass | Photo credit: E. A. Lazo-Wasem, Yale Peabody Museum, 2013.

The low marsh edge: cordgrass shaped by snails and crabs

At the daily-flooded edge of the marsh, smooth cordgrass (Spartina alterniflora) dominates. This narrow fringe marks the boundary between open water and marsh interior, where erosion pressure is highest and stability matters most.

Left: Healthy smooth cordgrass (Spartina alterniflora) line the estuary edge in Surf City, NC. | Photo credit: A. Mitchell, 2022. Right: Salt marsh die-off from grazing stress by marsh periwinkle snails and reduced predation by crabs, such as blue crabs, can create bare mudflats. | Photo credit: By Esuglia at English Wikipedia, CC BY-SA 3.0, https://commons.wikimedia.org/w/index.php?curid=65794096

Cordgrass growth here is tightly regulated by the marsh periwinkle snail (Littoraria irrorata). These snails climb grass stems to avoid inundation and graze directly on living tissue, often intensifying damage by facilitating fungal infection. At high densities, periwinkle grazing can dramatically reduce cordgrass height and biomass, effectively mowing the marsh edge (Silliman & Zieman, 2001).

Marsh periwinkle snails (Littoraria irrorata) are a common sight on cordgrass (Spartina alterniflora) in North Carolina | Photo credit: North Carolina Aquarium at Roanoke Island, 2018.

Unchecked grazing can destabilize the marsh platform—but periwinkles themselves are regulated by crabs, including blue crabs (Callinectes sapidus), fiddler crabs (Genus Uca), purple marsh crabs (Sesarma reticulatum), hermit crabs and other burrowing species. Crabs prey on snails, limiting grazing pressure and indirectly protecting cordgrass (Silliman et al., 2005).

Crabs also function as ecosystem engineers. Their burrows aerate sediments, relieve sulfide stress around plant roots, and improve tidal water movement through compacted soils (Bertness, 1985; Thomas & Blum, 2010). Where crabs are abundant, cordgrass grows taller and denser; where they are lost, marsh die-off can occur rapidly.

This zone persists through a trophic cascade: grass builds land, snails limit grass, and crabs keep the system in balance.

Mid-marsh: mussels and detritus processors reinforce the platform

Just upslope, where flooding becomes less frequent, plant communities shift toward mixtures that often include saltmeadow cordgrass (Spartina patens). Here, the ribbed mussel (Geukensia demissa) emerges as a key stabilizing force.

Saltmeadow cordgrass (Spartina patens) is an important marsh stabilizer that has higher productivity when it grows near ribbed mussel aggregations | Photo credit: Kristie Gianopulos

Ribbed mussels form dense clusters at the base of marsh vegetation, binding sediments with byssal threads and physically reinforcing marsh soils against erosion (Bertness, 1984). As filter-feeders, they concentrate nutrients by removing organic matter from tidal waters and depositing nitrogen-rich biodeposits directly into marsh sediments (Jordan & Valiela, 1982).

Ribbed mussels (Geukensia demissa) at the base of marsh vegetation | Photo credit: R. Bachand

Grasses growing near mussel aggregations exhibit higher productivity than those without mussels, demonstrating a strong facilitative relationship between animals and plants (Bertness, 1984). As vegetation senesces, detritivorous worms, insects, and microbial decomposers break down dead plant material, converting standing biomass into detritus that fuels food webs throughout the estuary (Mann, 1988).

The mid-marsh functions as a processing zone, reinforcing marsh structure while converting plant matter into usable energy.

High marsh: microbes that manage chemical stress

In the high marsh, dominated by black needlerush (Juncus roemerianus) and saltmeadow cordgrass (Spartina patens), flooding is limited to spring tides and storms. Prolonged exposure to air creates harsh soil conditions, including elevated salinity and sulfide accumulation.

Black needlerush grass (Juncus roemerianus) dominates the high marsh | Photo credit: ©Andy Newman

Here, microbial communities play a central role. Sulfate-reducing and sulfur-oxidizing bacteria regulate sulfide concentrations that would otherwise become toxic to plant roots, while microbial decomposition controls nutrient availability under fluctuating oxygen conditions (Howarth & Giblin, 1983).

Beneath the marsh surface, soil microbes regulate decomposition, carbon exchange, and chemical stress. Changes in salinity and flooding reshape microbial communities, influencing how marsh soils process organic matter and support vegetation across tidal elevations. | Image credit: Zhang et al., 2023.

Small soil invertebrates maintain pore spaces that allow brief pulses of oxygenated water to penetrate during flooding. Unlike the visibly engineered low marsh, the high marsh is stabilized largely through biogeochemical regulation rather than grazing or predation.

This zone endures because microbes quietly buffer plants against chemical extremes.

From microbes in the soil to grasses at the surface, biological interactions drive marsh formation. Microbial processes govern decomposition and organic matter buildup, helping determine whether marsh platforms gain elevation, remain stable, or collapse | Image credit: Abbot, Quirk & Fultz, 2022.

The marsh–upland transition: keeping the boundary intact

At the uppermost margin of the marsh, tidal influence becomes intermittent and environmental stress shifts from salinity to erosion and freshwater input. Burrowing invertebrates increase soil permeability, allowing stormwater and tidal surges to infiltrate rather than scour the surface (Thomas & Blum, 2010).

A profile illustration . depicting the recommended transition of plant types from the edge of the salt marsh to the upland buffer. | Image credit: Massachusetts Office of Coastal Zone Management

Vegetation root networks stabilize soils exposed to drying and wave action, while animal burrows act as pressure-release pathways during extreme events. When these biological processes are disrupted—by shoreline hardening or vegetation removal—the marsh edge often collapses abruptly rather than adjusting gradually.

This boundary holds only as long as water can move through it.

Black, organic-rich peat exposed after storms marks the remains of an ancient salt marsh once buried beneath barrier sands. Its reappearance along North Topsail Beach records long-term shoreline change and marsh migration. Photo credit: Bill Tresnan, 2024.

A marsh built by interactions

Across all elevations in Onslow County marshes, the pattern is consistent:

Plants define the zones—but animals and microbes determine whether those zones endure.

Conceptual diagram of revised juvenile blue crab ontogenetic habitat shifts. Arrows depict transitions between habitats with increases in size. Arrow widths denote abundance contributions of individuals between habitats. | Image credit: Hyman et al., 2023

From grazers that keep seagrass blades clean, to crabs that hold the marsh edge together, to microbes that manage invisible chemical stress, the marsh is sustained by small organisms with outsized influence. Together, these interactions determine not just what lives in the marsh, but whether the marsh itself endures.

Purple marsh crabs (Sesarma reticulatum) moving together along the marsh edge on South Topsail Island, North Carolina. Their collective movement and feeding activity illustrate how small organisms play outsized roles in maintaining marsh structure. Photo credit: A. Mitchell, 2025.

References

Abbott, K. M., Quirk, T., & Fultz, L. M. (2022). Soil microbial community development across a 32-year coastal wetland restoration time series and the relative importance of environmental factors. Science of The Total Environment, 821, 153359. https://doi.org/10.1016/j.scitotenv.2022.153359

Bertness, M. D. (1984). Ribbed mussels and Spartina Alterniflora production in a New England salt marsh. Ecology, 65(6), 1794-1807. https://doi.org/10.2307/1937776

Bertness, M. D. (1985). Fiddler crab regulation of Spartina alterniflora production on a New England salt marsh. Ecology, 66(3), 1042-1055. https://doi.org/10.2307/1940564

Duffy, J. E., Reynolds, P. L., Boström, C., Coyer, J. A., Cusson, M., Donadi, S., Douglass, J. G., Eklöf, J. S., Engelen, A. H., Eriksson, B. K., Fredriksen, S., Gamfeldt, L., Gustafsson, C., Hoarau, G., Hori, M., Hovel, K., Iken, K., Lefcheck, J. S., Moksnes, P., … Stachowicz, J. J. (2015). Biodiversity mediates top–down control in eelgrass ecosystems: A global comparative‐experimental approach. Ecology Letters, 18(7), 696-705. https://doi.org/10.1111/ele.12448

Howarth, R. W., & Giblin, A. (1983). Sulfate reduction in the salt marshes at Sapelo island, Georgia. Limnology and Oceanography, 28(1), 70-82. https://doi.org/10.4319/lo.1983.28.1.0070

Hyman, A. C., Chiu, G. S., Seebo, M. S., Smith, A., Saluta, G. G., Knick, K. E., & Lipcius, R. N. (2023). Model-based evaluation of critical nursery habitats for juvenile blue crabs through ontogeny: Abundance and survival in seagrass, salt marsh, and unstructured bottom. https://doi.org/10.1101/2023.07.20.549877

Jordan, T. E., & Valiela, I. (1982). A nitrogen budget of the ribbed mussel, Geukensia demissa, and its significance in nitrogen flow in a New England salt marsh. Limnology and Oceanography, 27(1), 75-90. https://doi.org/10.4319/lo.1982.27.1.0075

Mallin, M. A., Burkholder, J. M., Cahoon, L. B., & Posey, M. H. (2000). North and South Carolina coasts. Marine Pollution Bulletin, 41(1-6), 56-75. https://doi.org/10.1016/s0025-326x(00)00102-8

Mann, K. H. (1988). Production and use of detritus in various freshwater, estuarine, and coastal marine ecosystems. Limnology and Oceanography, 33(4part2), 910-930. https://doi.org/10.4319/lo.1988.33.4part2.0910

Orth, R. J., Heck, K. L., & Van Montfrans, J. (1984). Faunal communities in seagrass beds: A review of the influence of plant structure and prey characteristics on predator: Prey relationships. Estuaries, 7(4), 339. https://doi.org/10.2307/1351618

Orth, R. J., & Van Montfrans, J. (1984). Epiphyte-seagrass relationships with an emphasis on the role of micrograzing: A review. Aquatic Botany, 18(1-2), 43-69. https://doi.org/10.1016/0304-3770(84)90080-9

Silliman, B. R., Van de Koppel, J., Bertness, M. D., Stanton, L. E., & Mendelssohn, I. A. (2005). Drought, snails, and large-scale die-off of southern U.S. salt marshes. Science, 310(5755), 1803-1806. https://doi.org/10.1126/science.1118229

Silliman, B. R., & Zieman, J. C. (2001). Top-down control of Spartina alterniflora production by periwinkle grazing in a Virginia salt marsh. Ecology, 82(10), 2830. https://doi.org/10.2307/2679964

Thomas, C., & Blum, L. (2010). Importance of the fiddler crab Uca pugnax to salt marsh soil organic matter accumulation. Marine Ecology Progress Series, 414, 167-177. https://doi.org/10.3354/meps08708

Valentine, J. F., & Duffy, J. E. (n.d.). The central role of grazing in seagrass ecology. Seagrasses: Biology, Ecology and Conservation, 463-501. https://doi.org/10.1007/1-4020-2983-7_20

Zhang, G., Bai, J., Jia, J., Wang, W., Wang, D., Zhao, Q., Wang, C., & Chen, G. (2023). Soil microbial communities regulate the threshold effect of salinity stress on SOM decomposition in coastal salt marshes. Fundamental Research, 3(6), 868-879. https://doi.org/10.1016/j.fmre.2023.02.024

January 10, 2026
Foraminifera: The Marsh’s Memory Keepers

What microscopic shells along Topsail and Surf City tell us about ancient seas, living marshes, and the future coastline

On a winter walk along the marsh edge in Topsail or Surf City, the landscape feels quiet. Cordgrass has faded to straw, tidal creeks run clear, and storm tides have pulled back layers of sediment that were hidden just months ago. Winter slows the marsh, but it also reveals it. Along exposed creek banks and tidal flats, the smallest residents of these ecosystems leave behind subtle traces — grains, spirals, and pin-sized shells that most people would mistake for sand.

These are the remains of foraminifera, key marsh indicators, and they carry a record far older than the marsh itself (Murray, 2006; Scott et al., 2001).

What Are Foraminifera?

Foraminifera, often called forams, are single-celled marine organisms — not animals, but protists — that live in oceans, estuaries, and salt marshes around the world (Murray, 2006). Despite their microscopic size, most foraminifera build protective shells, known as tests, made either from calcium carbonate or from tiny grains of sediment cemented together (Scott et al., 2001; Debenay & Guillou, 2002).

Different species occupy very specific zones within a marsh. Some live high in the intertidal, others closer to open water. Their distribution reflects precise environmental conditions such as salinity, tidal elevation, oxygen availability, and sediment type (Edwards et al., 2004; Culver & Horton, 2005). Because of this tight ecological coupling, foraminifera respond quickly when conditions change (Debenay & Guillou, 2002).

Peneropolis proteus is the one of three most dominant species of fossil foraminifera in the Onslow Bay area, occurring in about 15% of samples (Schnitker, 1971).

Why Winter Reveals the Record

In summer, marsh surfaces are busy and obscured. Dense vegetation, algae, burrowing organisms, and constant sediment mixing make it difficult to see what lies beneath. In winter, vegetation thins, biological activity slows, and storm tides rework creek edges and tidal flats. Fine sediments are redistributed, exposing layers that formed years, decades, or even centuries earlier (Scott et al., 2001; Gehrels, 1994).

Winter does not create this record — it simply makes it visible (Murray, 2006).

Size, Stability, and Ancient Seas

Some fossil foraminifera grew to the size of coins, while most living forms today are no larger than grains of sand (Murray, 2006). This contrast reflects the environments they evolved within. In ancient shallow seas, conditions were often warm, stable, and chemically consistent for long periods of time. Temperature, salinity, and carbonate availability changed slowly, allowing foraminifera to grow over many years, build thick and complex shells, and, in some cases, form partnerships with symbiotic algae — similar to the relationship between corals and the algae that live within their tissues — which provided an additional energy source through photosynthesis (Hallock, 1981; Murray, 2006). These systems favored persistence and size.

Over time, coastlines shifted and sea levels changed, giving rise to the highly dynamic estuaries and marshes we see today. In these modern environments, conditions can fluctuate over hours or seasons. Salinity rises and falls, oxygen levels vary, sediments are rearranged, and water chemistry responds quickly to storms and freshwater input (Debenay & Guillou, 2002; Culver & Horton, 2005). Under such variability, smaller foraminifera that grow rapidly and tolerate change are more likely to survive. Because foraminifera respond directly to these environmental conditions, even subtle shifts can reorganize their communities, altering shell size, composition, and diversity in ways that can persist in sediments long after the initial change has occurred (Edwards et al., 2004; Kemp et al., 2013).

Tiny Shells, Deep Time: How Marshes Remember

Foraminifera are among the most powerful tools scientists use to reconstruct ancient coastal ecosystems because the conditions they live in are permanently recorded in their shells. Individual species occupy narrow ecological ranges defined by salinity, tidal elevation, oxygen availability, temperature, and sediment type. Because of this specificity, the particular mix of foraminifera preserved in a layer of marsh sediment reflects the environmental conditions present when that layer formed.

When scientists extract sediment cores from marshes, they are not looking for isolated snapshots in time, but for transitions. As layers accumulate, changes in species composition, shifts between calcium-based shells and sediment-built shells, and variations in diversity reveal how marsh conditions evolved. These biological signals can indicate changes in flooding frequency, sediment stability, freshwater influence, and tidal reach — often aligning with known shifts in sea level or shoreline position.

What makes foraminifera especially valuable is that they record change continuously. Each generation reflects the conditions it experienced, leaving behind a layered biological archive that links past marshes to present ones — comparable to how sedimentary layers exposed in the Grand Canyon record changing environments over deep time.This continuity allows scientists to distinguish gradual environmental adjustment from more abrupt change and to assess whether modern conditions resemble states marshes have previously endured — or represent departures from historical patterns.

Quinqueloculina seminula (left) and Plancopsilina confusa (right) are the top three most dominant species of fossil foraminifera in the Onslow Bay area, each occurring in about 20% of samples (Schnitker, 1971).

What Lives in a Handful of Marsh Sand

If you scoop a small handful of sand or mud from a North Carolina marsh and let it dry, it looks ordinary—grains, bits of plant matter, flecks of shell. Where sediment cores reveal depth at the scale of decades and centuries, living marsh surfaces show that same pattern compressed into just a few centimeters. But research from the Outer Banks suggests that even this unremarkable material holds a surprisingly rich living community.

Foraminifera under biological microscope with sand.

In a detailed study of marsh sediments along the North Carolina coast, scientists examined not just which foraminifera were present, but which ones were alive at the time of sampling. What they found was not a thin layer of life resting at the surface, but a vertically structured community extending down into the sediment itself (Culver, 2005).

Some foraminifera lived right at the surface, where tides regularly wash over the marsh. Others occupied sediments a centimeter or more below, in darker, less oxygenated layers. In total, more than twenty species were documented living within marsh sediments, their distributions shaped by subtle differences in tidal flooding, salinity, and marsh elevation (Culver, 2005).

Not all species were equally widespread. A few, including Jadammina macrescens and Tiphotrocha comprimata, appeared across multiple sites and depths, suggesting a tolerance for changing marsh conditions. Many others were more selective, occurring only in certain zones or at particular depths. This means that even small changes in where you stand—closer to a tidal creek or higher on the marsh platform—can correspond to a different microscopic community beneath your feet (Culver, 2005).

Upper image: Jadammina macrescens under microscope.| Image credit: Parker, G. G., Phleger, et al. 1953. Cushman Found.Foram.Research Spec.Pub. (n.2): 15, pl.3,f.8.
Lower image: Tiphotrocha comprimata under microscope | Image credit: Hesemann, M., The Foraminifera.eu Database (2026). Accessed at http://www.foraminifera.eu.
https://doi.org/10.13140/RG.2.2.22727.11680/1.

As these organisms die, their shells remain. Layer by layer, those shells become part of the sediment, preserving a record of where tides reached, how often flooding occurred, and how stable the marsh surface was at that moment in time (Scott et al., 2001). What begins as a living community quietly becomes part of the marsh’s long-term record.

Although the Outer Banks are not identical to the marshes behind Topsail and Surf City, the pattern holds across North Carolina’s coast: foraminifera respond to local conditions at very small scales. Their presence, abundance, and depth within the sediment shift from place to place, reflecting the marsh’s relationship with water, salt, and time (Edwards et al., 2004; Culver & Horton, 2005).

Cibicidoides bradyi (horizontal scale bar = 200μm, vertical scale bar = 400μm) occur in less than 20 m at about 1% of samples in the Onslow County area (Schnitker, 1971).

For someone walking the marsh in winter, this means that the sand exposed along a creek bank carries more than the imprint of the last storm. It carries traces of countless tides before it—each one leaving behind shells small enough to escape notice, yet durable enough to remember.

What Changes in Foraminifera Mean for the Ecosystem

An example of how shifts in reef communities reflect shifts in foraminiferal communities below (Prazeres, Martínez-Colón & Hallock, 2020).

Foraminifera do not exist in isolation. They are part of the marsh food web, contributing to the transfer of energy and nutrients from microscopic primary producers to larger organisms (Murray, 2006). Many small invertebrates consume foraminifera directly, while others rely on the microbial communities and organic matter associated with their shells (Debenay & Guillou, 2002). In turn, these invertebrates support fish, crabs, and birds that depend on marsh productivity (Scott et al., 2001).

When foraminiferal communities shift, the effects can ripple outward. A decline in diversity or a move toward stress-tolerant species often reflects changes in sediment stability, oxygen availability, or salinity — conditions that also influence marsh plants, benthic invertebrates, and juvenile fish habitat (Culver & Horton, 2005; Edwards et al., 2004). In this way, changes in foraminifera can foreshadow broader ecological adjustments, even when the marsh surface still appears healthy (Debenay & Guillou, 2002).

Because foraminifera respond quickly to environmental change, they often register these shifts before larger organisms do. Their shells capture early signals of altered flooding patterns, reduced sediment input, or changing water chemistry (Gehrels, 1994; Kemp et al., 2013). What follows may be changes in plant community structure, altered nutrient cycling, or shifts in the species that use marshes as nursery grounds. Foraminifera do not cause these changes, but they reveal when the system’s internal balance begins to shift (Scott et al., 2001).

Reading Change in Living Marshes

Salt marshes are dynamic systems by nature. They grow, erode, migrate, and rebuild as sediment moves and sea level changes (Kemp et al., 2013). The challenge for scientists is distinguishing normal variability from directional change — shifts that push marshes beyond the conditions they have historically been able to tolerate. Foraminifera are especially useful in making that distinction because they respond quickly and directly to their surroundings (Debenay & Guillou, 2002).

When marsh conditions move outside typical ranges — whether through altered hydrology, changes in sediment supply, or shifts in salinity — foraminiferal communities reorganize. Species diversity may decline, stress-tolerant forms can become dominant, and assemblages tied to specific tidal elevations may disappear (Culver & Horton, 2005). These changes often occur before larger, more visible signs of stress appear, such as widespread plant die-off or shoreline erosion (Edwards et al., 2004). In this sense, foraminifera act as early responders, recording change while the marsh still appears intact at the surface (Scott et al., 2001).

Along the marshes behind Topsail and Surf City, this sensitivity gives foraminifera particular importance. They help establish local baselines for what healthy marsh conditions look like, provide context for interpreting present-day shifts, and preserve a record of the conditions that supported marsh stability in the past (Culver & Horton, 2005; Kemp et al., 2013). By linking modern observations to sedimentary records, foraminifera allow scientists to ask not only what is changing, but how quickly change is occurring and whether it remains within the range marshes have previously endured. Understanding marsh resilience in this way is not abstract or theoretical — it is grounded in the specific history and behavior of this coastline.

Salt marsh in Surf City, NC. | Photo credit: Mitchell (2026)

Closing

Standing at the marsh edge in winter, it is easy to miss the smallest details. Yet beneath the quiet surface, microscopic shells record centuries of change — how water moved, how shorelines shifted, and how marshes adapted (Murray, 2006). Foraminifera remind us that long before satellites or tide gauges, coastlines were already keeping their own records. All we have to do is learn how to read them.

References

Culver, S. J. (2005). Infaunal marsh foraminifera from the Outer Banks, North Carolina, U.S.A. The Journal of Foraminiferal Research, 35(2), 148-170. https://doi.org/10.2113/35.2.148

Debenay, J., & Guillou, J. (2002). Ecological transitions indicated by foraminiferal assemblages in paralic environments. Estuaries, 25(6), 1107-1120. https://doi.org/10.1007/bf02692208

Edwards, R., Wright, A., & Van de Plassche, O. (2004). Surface distributions of salt-marsh foraminifera from Connecticut, USA: Modern analogues for high-resolution sea level studies. Marine Micropaleontology, 51(1-2), 1-21. https://doi.org/10.1016/j.marmicro.2003.08.002

Gehrels, W. R., & Kemp, A. C. (2021). Salt marsh sediments as recorders of Holocene relative sea-level change. Salt Marshes, 225-256. https://doi.org/10.1017/9781316888933.011

Hallock, P. (1981). Algal symbiosis: A mathematical analysis. Marine Biology, 62(4), 249-255. https://doi.org/10.1007/bf00397691

Kemp, A. C., Horton, B. P., Vane, C. H., Berhhardt, C. E., Corbett, D. R., Engelhart, S. E., Anisfeld, S. C., Parnell, A. C., & Cahill, N. (2013). Sea-level change during the last 2500 years in New Jersey, USA. Quaternary Science Reviews, 81(2013), 90-104. https://www.whoi.edu/cms/files/Kemp2013QSR_170144.pdf

Murray, J. W. (2006). Ecology and applications of benthic foraminifera. Cambridge University Press.

Schnitker, D. (1971). Distribution of Foraminifera on the North Carolina Continental Shelf. Tulane Studies in Geology and Paleontology, 8(4), 169-215. https://journals.tulane.edu/tsgp/article/view/560

Scott, D. B., Medioli, F. S., & Schafer, C. T. (2001). Monitoring in coastal environments using foraminifera and Thecamoebian indicators. Cambridge University Press.

January 3, 2026