T
Peltier, W. R., 1998. Postglacial variations in the level of the sea:
TECTONIC EUSTASY
Nils-Axel Mörner
Paleogeophysics and Geodynamics, Saltsjöbaden,
Sweden
Synonyms
Tectono-eustasy
Definition
Tectonic eustasy (often written, tectono-eustasy) denotes
the sea-level changes generated by changes in ocean
basin volume (e.g., Mörner, 1986; Mörner, 1987; Rona,
1995).
Description
Changes in ocean basin volume are generated by different
geodynamic processes such as sea-floor subsidence, ridge
growth, plate motions, and crustal movements. It is a slow
process leading to maximum sea-level changes of

0.06 mm/year (Mörner, 1996). Global isostatic adjust-
ment (GIA) in response to the glacial loading/de-loading
in association with ice ages (e.g., Peltier, 1982, 1998)
may be considered a special element of tectonic eustasy.
Bibliography
Mörner, N.-A., 1986. The concept of eustasy: a redefinition. Jour-
nal of Coastal Research, Special Issue 1, 49–51.
Mörner, N.-A., 1987. Models of global sea level changes. In Tooley,
M. J., and Shennan, I. (eds.), Sea Level Changes. Hoboken:
Blackwell, pp. 333–355.
Mörner, N.-A., 1996. Rapid changes in coastal sea level. Journal of
Coastal Research, 12, 797–800.
Peltier, W. R., 1982. Dynamics of the ice age Earth. Advances of
Geophysics, 24, 1–144.
M.J. Kennish (ed.), Encyclopedia of Estuaries, DOI 10.1007/978-94-017-8801-4,
© Springer Science+Business Media Dordrecht 2016
implications for climate dynamics and solid-earth geophysics.
Reviews of Geophysics, 38, 603–689.
Rona, P. A., 1995. Tectonoeustasy and Phanerozoic sea levels. In
Finkl Jr., C. W. (ed.), Holocene Cycles: Climate, Sea Levels,
and Sedimentation. Journal of Coastal Research, Special Issue
17, 269–277.
Cross-references
Shoreline Changes
Submerged Coasts
Submergent Shoreline
THERMAL BIOLOGY
Nicholas B. Colvard and Brian Helmuth
Marine Science Center, Northeastern University,
Nahant, MA, USA
Synonyms
Biophysical ecology; Organism temperature; Thermal
physiology
Definition
Thermal biology is the study of physiological and ecolog-
ical consequences of body temperature and of the
biophysical, morphological, and behavioral determinants
of organism temperature.
Introduction
The temperature of an organism’s body affects almost all
physiological processes, which in turn can have signifi-
cant consequences for organisms, and interactions such
as competition and predation. While some animals (and
a few plants) make significant amounts of metabolic heat
662 THERMAL BIOLOGY

Thermal Biology, Figure 1 Photographic image of the intertidal community (left) demonstrates species vertical zonation and substrate
heterogeneity, with concurrent infrared (IR) thermal image (right) of the same site during low tide. The IR image demonstrates the
large variability in temperatures (15–37  C) observed in this small section of shore, which may in part drive small-scale distribution and
zonation patterns. Photograph courtesy of N. Colvard and B. Helmuth, FLIR E60 Infrared Camera.

(i.e., are endothermic), the body temperatures of estuarine surfaces lose considerable amounts of heat via the evapo-
invertebrates and plants are driven by the exchange of heat ration of water, although this water loss potentially comes
with the ambient environment – they are ectothermic – at the cost of desiccation (Iacarella and Helmuth, 2011).
and can vary substantially over the course of a day, season, As above, the movement of organisms into shaded crev-
and year. ices, on the shaded sides of rocks, or depth in the sediment
can lead to temperature differences over the scale of centi-
meters that exceed those observed over thousands of kilo-
Heat exchange and body temperature meters of latitude (Denny et al., 2011).
Intertidal organisms in estuaries are subject to diel tidal fluc-
tuations, and so their body temperatures at low tide aerial Direct and indirect effects of body
exposure can significantly differ than that during submer- temperature on estuarine organisms
gence (Jost and Helmuth, 2007). The dominant driver
Organism temperature has numerous effects on physiolog-
of heat input in these systems is solar radiation, so that the
ical processes in estuarine organisms (Somero, 2011),
temperature of an animal’s or a plant’s body is generally
including functioning of critical enzymes and other
much warmer than the temperature of the surrounding air
cellular functions, changes in metabolic rate, and increases
during the day; at night, as heat is lost through infrared
in metabolic oxygen demand (Pörtner, 2010). For an
radiation, organisms can be several degrees cooler than
excellent overview of physiological impacts of tempera-
the air. The amount of shading can have a large influence
ture on marine organisms, see Somero (2011). The thermal
as can the amount of wind (convection). Thus, for example,
biology of estuarine organisms also indirectly affects spe-
snails at the top of a marsh canopy can have temperatures
cies through interactions such as competition and preda-
different from animals near the substrate (Iacarella and
tion (Russell et al., 2012). Wethey (1984) showed that
Helmuth, 2011). Solar heating can also increase the temper-
the competitive ability of barnacles is affected by their
ature of shallow waters so that continually submerged estu-
temperatures and that the zonation of two barnacle species
arine organisms can in some cases experience large
could be explained by their differential physiological sen-
fluctuations in temperature.
sitivity to body temperature. Sanford (2002) demonstrated
The behavioral and morphological characteristics of an
that rates of predation by the keystone sea star Pisaster
organism dictate the rate of heat transfer between the sur-
were elevated at increased water temperature, and con-
rounding environment and the organism, and the size,
versely, Pincebourde et al. (2008) reported decreases in
shape, mass, and color of an organism can affect body
predation rate by up to 40 % during elevated temperatures
temperature. Thus, two organisms exposed to identical
at low tide in air.
environmental conditions can display very different body
temperatures from one another, even when neither is mak-
ing any appreciable metabolic heat (Figure 1). The behav- Monitoring temperature
ior of an organism can also affect rates of heating and In water, the temperature of a marine organism approxi-
cooling. Organisms such as algae and sea stars with wet mates that of the surrounding water, although in some
 THERMAL BIOLOGY
cases of low water flow and high solar radiation, tempera-
tures of organisms such as corals can be several degrees
warmer than the water around them (Jimenez et al.,
2008). Gradients in water temperature in many coastal
systems can be very steep, so that the temperature even
a few meters below the surface can be considerably cooler
than that recorded on the surface, for example, by satellite
(Castillo and Lima, 2010). Measuring the temperatures of
organisms in situ and at scales appropriate to the organism
in question is therefore vital for understanding their ther-
mal biology.
During aerial exposure at low tide, animal and plant
temperatures are often quite different from those of the
surrounding air and substrata (Lathlean et al., 2011). Frus-
tratingly, the same factors that lead to observed differences
in organism temperature also affect the temperature
recorded by instruments, such as now commonly used
data loggers, so that the size, color, and mass of the instru-
ment (or worse yet, the casing in which it is housed) all
affect the temperature that it records. Fitzhenry et al. (2004)
reported average differences of 14  C between the body
temperature of intertidal mussels and the adjacent temper-
ature loggers. To combat this issue, the use of biomimetic
temperature loggers has been adopted (Lima et al., 2011).
Biomimetics are “thermally matched” to organisms and
can in many cases record temperatures close to that of
study organisms (Lima et al., 2011). Notably, a central
message is that a single instrument almost never records
measurements that are applicable to all organisms, since
each can display a different temperature.
The use of infrared thermography (Figure 1) has also
been used to measure patterns of temperature in the field
(Lathlean et al., 2012), although this approach carries with
it some drawbacks as well, as the camera needs to be cal-
ibrated to the surface properties (emissivity) of each spe-
cies and the instrument cannot be deployed continuously
for long time periods.
Climate change and thermal biology
The relevance of thermal biology in coastal regions has
received renewed attention in the face of climate change.
Changes in the distribution and abundance of many
coastal species have been reported (Southward
et al., 2005). These studies have emphasized that if we
are to predict the likely impacts of ongoing and projected
climate change on coastal species, a nested approach that
considers the physiological, ecological, and biogeo-
graphic mechanisms by which the environment affects
the thermal biology of organisms in what are likely to be
unprecedented conditions should be used (Howard
et al., 2013).
Summary
The study of thermal biology in estuarine species is very
important since these organisms are subject to daily fluctu-
ations in environmental conditions from ebb and flood
tides. Estuarine organisms can serve as model systems
663
when evaluating the impact of climate change and associ-
ated environmental processes because of the unique inter-
change with terrestrial and aquatic environments. In some
localities estuarine organisms are living at their thermal
maximum, and with the ensuing threat of aerial and water
temperature increases in the coming century, it will be par-
amount to understand how these changes will impact the
primary producers and ecosystem engineers in estuarine
biotic communities.
Bibliography
Castillo, K. D., and Lima, F. P., 2010. Comparison of in situ and
satellite-derived (MODIS-Aqua/Terra) methods for assessing
temperatures on coral reefs. Limnology and Oceanography:
Methods, 8, 107–117.
Denny, M. W., Dowd, W. W., Bilir, L., and Mach, K. J., 2011.
Spreading the risk: small-scale body temperature variation
among intertidal organisms and its implications for species per-
sistence. Journal of Experimental Marine Biology and Ecology,
400, 175–190.
Fitzhenry, T., Halpin, P. M., and Helmuth, B., 2004. Testing the
effects of wave exposure, site, and behavior on intertidal mussel
body temperatures: applications and limits of temperature logger
design. Marine Biology, 145, 339–349.
Howard, J., Babij, E., Griffis, R., Helmuth, B., Himes-Cornell, A.,
Neimier, P., Orbach, M., Petes, L., Allen, S., Auad, G., Beard,
R., Boatman, M., Bond, N., Boyer, T., Brown, D., Clay, P.,
Crane, K., Cross, S., Dalton, M., Diamond, J., Diaz, R., Dortch,
Q., Duffy, E., Fauquier, D., Fisher, W., Graham, M., Halpern, B.,
Hansen, L., Hayum, B., Herrick, S., Hollowed, A., Hutchins, D.,
Jewett, E., Jin, D., Knowlton, N., Kotowicz, D., Kristiansen, T.,
Little, P., Lopez, C., Loring, P., Lumpkin, R., Mace, A.,
Mengerink, K., Morrison, J. R., Murray, J., Norman, K.,
O’Donnell, J., Overland, J., Parsons, R., Pettigrew, N., Pfeiffer,
L., Pidgeon, E., Plummer, M., Polovina, J., Quintrell, J., Rowles,
T., Runge, J., Rust, M., Sanford, E., Send, U., Singer, M., Speir,
C., Stanitski, D., Thornber, C., and Xue, Y., 2013. Oceans and
marine resources in a changing climate. Oceanography and
Marine Biology Annual Review, London, 51, 71–192.
Iacarella, J. C., and Helmuth, B., 2011. Body temperature and des-
iccation constrain the activity of Littoraria irrorata within the
Spartina alterniflora canopy. Journal of Thermal Biology, 37,
15–22.
Jimenez, I. M., Kühl, M., Larkum, A. W. D., and Ralph, P. J., 2008.
Heat budget and thermal microenvironment of shallow-water
corals: do massive corals get warmer than branching corals?
Limnology and Oceanography, 53, 1548–1561.
Jost, J., and Helmuth, B., 2007. Morphological and ecological deter-
minants of body temperature of Geukensia demissa, the Atlantic
ribbed Mussel, and their effects on mussel mortality. Biological
Bulletin, 213, 141–151.
Lathlean, J. A., Ayre, D. J., and Minchinton, T. E., 2011. Rocky
intertidal temperature variability along the southeast coast of
Australia: comparing data from in situ loggers, satellite-derived
SST and terrestrial weather stations. Marine Ecology Progress
Series, 439, 83–122.
Lathlean, J. A., Ayre, D. J., and Minchinton, T. E., 2012. Using
infrared imagery to test for quadrat-level temperature variation
and effects on the early life history of a rocky-shore barnacle.
Limnology and Oceanography, 57, 1279–1291.
Lima, F. P., Burnett, N. P., Helmuth, B., Aveni-Deforge, K., Kish,
N., and Wethey, D. S., 2011. Monitoring the intertidal environ-
ment with bio-mimetic devices. In Biomimetic Based Applica-
tions. Rijeka: INTECH Publishing, pp. 499–522.
664 TIDAL ASYMMETRY
Pincebourde, S., Sanford, E., and Helmuth, B., 2008. Body temper-
ature during low tide alters the feeding performance of a top
intertidal predator. Limnology and Oceanography, 53,
1562–1573.
Pörtner, H. O., 2010. Oxygen- and capacity-limitation of thermal
tolerance: a matrix for integrating climate-related stressor effects
in marine ecosystems. Journal of Experimental Biology, 213,
881–893.
Russell, B. D., Harley, C. D. G., Wernberg, T., Mieszkowska, N.,
Widdicombe, S., Hall-Spencer, J. M., and Connell, S. D., 2012.
Predicting ecosystem shifts requires new approaches that inte-
grate the effects of climate change across entire systems. Biology
Letters, 8, 164–166.
Sanford, E., 2002. Water temperature, predation, and the neglected
role of physiological rate effects in rocky intertidal communities.
Integrative and Comparative Biology, 42, 881–891.
Somero, G. N., 2011. Comparative physiology: a “crystal ball” for
predicting consequences of global change. American Journal
of Physiology- Regulatory Integrative and Comparative Physi-
ology, 301, R1–R14.
Southward, A. J., Langmead, O., Hardman-Mountford, N. J.,
Aitken, J., Boalch, G. T., Dando, P. R., Genner, M. J., Joint, I.,
Kendall, M. A., Halliday, N. C., Harris, R. P., Leaper, R.,
Mieszkowska, N., Pingree, R. D., Richardson, A. J., Sims,
D. W., Smith, T., Walne, A. W., Hawkins, S. J., et al., 2005.
Long-term oceanographic and ecological research in the western
English Channel. Advances in Marine Biology, 47, 1–105.
Wethey, D. S., 1984. Sun and shade mediate competition in the
barnacles Chthamalus and Semibalanus: a field experiment.
Biological Bulletin, 167, 176–185.
Cross-references
Climate Change
Ecological Niche
Environmental Gradients
Tides
TIDAL ASYMMETRY
Geórgenes H. Cavalcante
Institute of Atmospheric Science, Federal University of
Alagoas, Maceió, Alagoas, Brazil
Definition
Tidal asymmetry is defined by the discrepancies in the
duration of the falling and rising tides of water elevation.
This duration asymmetry may be manifested as an
inequality in flood/ebb tidal current magnitudes.
Description
Tidal asymmetry can produce two types of estuarine sys-
tems: flood dominant or ebb dominant. The system
referred to as flood dominant occurs when the duration
of the falling tide exceeds that of the rising tide leading
to a larger peak flood current (Headquarters, U.S. Army
Corps of Engineers, 2002); when the duration of the fall-
ing tide is shorter than that of the rising tide, it leads to
a stronger peak ebb current, and then the system is referred
to as ebb dominant.
The sources behind the asymmetrical tides are the inter-
actions of multiple tidal constituents (Boon and Byrne,
1981). Phase difference between constituents dictates the
direction of asymmetry (i.e., flood or ebb dominance),
while the ratio of constituent amplitudes reflects the
degree of distortion (Friedrichs and Aubrey, 1988).
Bibliography
Boon, J. D., and Byrne, R. J., 1981. On basin hypsometry and the
morphodynamic response of coastal inlet systems. Marine Geol-
ogy, 40(1–2), 27–48.
Friedrichs, C. T., and Aubrey, D. G., 1988. Non-linear tidal distor-
tion in shallow well-mixed estuaries: a synthesis. Estuarine
Coastal Shelf Science, 27(5), 521–545.
Headquarters, U.S. Army Corps of Engineers, 2002. Tidal Velocity
Asymmetry at Inlets. Washington, DC: ERDC/CHL CHETN-
IV-47.
Cross-references
Tidal Hydrodynamics
Tidal Ranges
TIDAL DATUM
Ivan D. Haigh
Ocean and Earth Science, National Oceanography Centre,
University of Southampton, European Way,
Southampton, UK
Definition
A tidal datum is a reference elevation of sea level, defined
in terms of a certain phase of a tide. Tidal datums are
primarily used to estimate water heights and/or depths
and determine horizontal boundaries. Tidal datums are
fundamental to the determination of spatial coordinates
of latitude, longitude, and elevation relative to mean sea
level. They are also essential as legal entities to establish
the limits of state jurisdiction over maritime zones
(NOAA, 2000).
Description
Tidal datums are local because they are estimated from
water level observations made at a particular tide gauge
site. Tidal ranges vary considerably spatially depending
on local hydrodynamic and topographic characteristics
(e.g., tidal range may be much larger at the head of an estu-
ary compared to the mouth), and hence tidal datums vary
accordingly. Therefore they should not be extended into
areas with significantly different oceanographic character-
istics without sufficient observations (or numerical model-
ing) to support any such extrapolation.
Tidal datums are usually estimated statistically from
water level observations made over a tidal datum epoch,
which is the interval of time recommended to calculate
tidal datums. This interval is usually at least 19 years, to
 TIDAL FLAT
ensure the full 18.6-year lunar nodal cycle is included in
the analysis. The tidal datum is the average water level
over a tidal epoch of water height at a particular tidal
phase.
A multitude of tidal datums are currently in use within
different counties. The common tidal datums are:
• Highest astronomical tide (HAT): The highest water
level predicted to occur under any combination of astro-
nomical conditions.
• Mean high water springs (MHWS) and mean low water
neaps (MHWN): The average of all high waters at the
time of spring or neap tide, respectively. This is applica-
ble only in regions with semidiurnal tides. In mixed and
diurnal tidal regions, mean higher high water (MHHW)
and mean lower high water (MLHW) are used, which
are the mean of the higher or lower of the two daily high
waters, respectively.
• Mean high water (MHW): The average of all high
waters.
• Mean sea level (MSL): The arithmetic mean of hourly
observations of water level.
• Mean low water springs (MLWS) and mean low water
neaps (MLWN): The average of all low waters at the
time of spring or neap tide, respectively. In mixed and
diurnal tidal regions, mean higher low water (MHLW)
and mean lower low water (MLLW) are used, which
are the mean of the higher or lower of the two daily
low waters, respectively.
• Mean low water (MLW): The average of all low waters.
• Lowest astronomical tide (LAT): The lowest water level
which can be predicted to occur under any combination
of astronomical conditions. LAT is often used to define
chart datum (CD), which is the water level which depths
displayed on a nautical chart are referenced to.
Bibliography
National Oceanic and Atmospheric Administration, 2000. Tidal
datums and their applications. NOAA Special Publication NOS
CO-OPS 1. Available from: http://tidesandcurrents.noaa.gov/
publications/tidal_datums_and_their_applications.pdf.
Cross-references
Mean Sea Level
Tidal Ranges
Tides
TIDAL FLAT
Vic Semeniuk
V & C Semeniuk Research Group, Warwick,
WA, Australia
Synonyms
Mud flat; Sand flat
665
Definition
A tidal flat is a low-gradient surface, underlain by mud,
muddy sand, or sand, which is exposed during low tide.
Definitional limits, zones, and characteristics
of tidal flats
Before ecological aspects of tidal flats are described, some
background information on the variety of tidal flats is pro-
vided from the perspective of their limits, zones, geomor-
phology, sedimentology, hydrology, and salinity, as these
will determine, for a given climate, the types of habitats
developed, the biota inhabiting them, and ecological pro-
cesses that will operate therein.
The concept and definition of a tidal flat varies with
author and between disciplines. Tidal flats, in the broadest
sense, are tidally exposed low-gradient surfaces, underlain
by a variety of substrates, and variably vegetated depending
on climate and location of the surface relative to mean sea
level (Semeniuk, 2005). They can be free of vegetation or
inhabited by mangroves, saltmarsh, or seagrass. In the
narrowest sense, for some authors, tidal flats are the
vegetation-free tidally exposed low-gradient surfaces usu-
ally occurring between MSL and low tide though, even in
this case, these types of flats may have a sparse ground
cover of seagrass. Some authors may incorrectly exclude
flats vegetated by mangroves (mangal) and saltmarsh, but
the vegetated parts of tidally exposed low-gradient surfaces
are still tidal flats. Although they have the same geomorphic
expression as traditional tidal flats, extensively exposed
rock pavements cut to low-gradient flats in the intertidal
zone, and gravel-floored flats are generally not considered
to be tidal flats. The consensus on tidal flats is that the tidally
exposed surface is underlain by soft sediment (sand, muddy
sand, or mud).
The moon and sun, the rotation of the earth, the regional
bathymetry, and the shape of the coast are major factors in
determining tidal hydrodynamics, the magnitude of tides,
and the forcing of the ebb and flow of tides (Cartwright,
1999). Water levels also can be affected by barometric
pressure and wind. The major effect of the moon and the
sun is to generate spring tides (the larger tidal oscillations
approximately coinciding with a new moon and full moon
or sun-earth-moon syzygy) and neap tides (the smaller
tidal oscillations approximately coinciding with the first
quarter moon and third quarter moon). Tidal zones
are identified as follows: equinoctial low water spring tide
(¼ELWS), a spring tide low water occurring twice yearly
during the equinoctial period; mean low water spring tide
(¼MLWS), the mean position of low water of spring tides;
mean low water neap tide (¼MLWN), the mean position
of low water of neap tides; MSL, the mean position of
water level between low water and high water; mean high
water neap tide (¼MHWN), the mean position of high
water during neap tides; mean high water spring tide
(¼MHWS), the mean position of high water during spring
tides; and equinoctial high water spring tide (¼EHWS),
a spring tide high water occurring twice yearly during
666
Tidal Flat, Figure 1 Tidal flat in profile showing the various tidal levels. Located between the supratidal and subtidal zones, the tidal
flat has the following levels: the lowest tidal level at equinoctial low-water spring tide (ELWS), mean low-water spring tide (MLWS),
mean low-water neap tide (MLN), mean sea level (MSL), mean high-water neap tide (MHN), mean high-water spring tide (MHWS), and
the highest tide level at equinoctial high-water spring tide (EHWS).
the equinoctial period. These various levels are shown dia-
grammatically in Figure 1. In this context, the entire sur-
face between ELWS and EHWS is the tidal zone, and
any low-gradient surfaces between these levels are tidal
flats. Above EHWS is the supratidal zone, and below
ELWS is the subtidal zone. During and the 1960s 1970s,
geological papers describing tidal flats in some regions
incorrectly termed the spring-tide-flooded surface
between MHWS and EHWS that dried out during the neap
tides as the “supratidal” zone. It should be noted that the
tidal zones delineated by tidal levels on a tidal flat may
not necessarily encompass the entire tidal interval (i.e.,
the extent of a tidal flat may not equate to the full extent
of the tidal zone) and the tidal flat may be confined, for
instance, to the interval between ELWS and MSL, or
EHWS and MHWN. Tidal intervals that are inhabited by
mangroves, saltmarsh, or seagrass or that are vegetation-
free are shown diagrammatically in Figure 2.
Tidal flats can directly adjoin a hinterland upland (and
be bordered along their landward edge by a cliff, by
a relatively more steeply sloping supratidal shore, or by
a dune) or adjoin a spit, sand bar, chenier, or sandy beach.
Rather than always being a simple uniformly sloping sur-
face from high water to low water, tidal flats often have
variation in gradients across the tidal interval (such as
low hummocks of sand, meters in width, on a mud flat,
or a low-relief chenier), as well as variation in substrate
types. The variation in slope is related to tidal level, tide
and wave energy gradients, and vegetation. Where there
is a range of sediment grain sizes from mud to sand on
the tidal flat, the dominant grain size, degree of exposure
to low-amplitude wave action when inundated, and the
tidal range, which determines current energy, will deter-
mine whether tidal flats are narrow or wide and whether
they are wholly sandy, wholly muddy, grade from sand
to mud from low tidal levels to high tidal levels, or the
reverse from mud to sand from low to high tidal level
TIDAL FLAT
(Figure 3). The frequency of inundation and the magni-
tude of tides determine the magnitude of tidal currents
and, hence, substrate type. For example, microtidal areas
generally have low-velocity tidal currents, while
macrotidal areas generate large currents. Further, with
wave action, mainly on sandy tidal flats, the tidal flat can
be shaped into shore-parallel, low-relief linear shoals
(or bars) and depressions, or low-relief shore-normal sand
bars. As a result of tidal current erosion, the tidal flat can
be traversed by dendritic to meandering shore-normal
tidal creeks. These smaller-scale variations influence the
distribution of biota across the tidal flat and the develop-
ment of small-scale habitats. Vegetation such as saltmarsh
and mangrove, in trapping sediment, can alter the slope of
the tidal flat in that there is a relative buildup of sediment
in the vegetated zone.
During low tide, many sandy tidal flats are covered by
a very thin film of mud as there generally is a small pro-
portion of mud in suspension in the waters flooding
a tidal flat which will settle, giving the impression that
the tidal flat is underlain by mud. While there are true
mud flats, many tidal flats that have been termed “mud
flats” actually are sandy tidal flats with a thin film of mud.
When tidal flats are exposed during low tide, water
under, or within a flat is subject to evaporation and transpi-
ration resulting in salinization, and a tidal flat salinity gradi-
ent is developed. This is systematic, progressive increase in
salinity in groundwater (as pore water) and pellicular water
(the water circumferential to sediment grains in a temporary
vadose zone) across the tidal flat. The salinity gradient is
described by Semeniuk (this volume) in “Tidal flat salinity
gradients.” The change in salinity across a tidal flat is due to
the interacting effects of inundation, evaporation, transpira-
tion, and sediment types. The salinity of groundwater and
pellicular water varies spatially and temporally depending
on (1) the position of a tidal flat in an estuary (and hence
the salinity of the recharging source water); tidal flats
 TIDAL FLAT 667

Tidal Flat, Figure 2 Idealized diagram of tidal flat surfaces showing a range of vegetation and plant life that may occupy specific tidal
zones. The positions of the various tide levels for these profiles are shown in Figure 1. (a) Saltmarsh on the high tidal flats usually
between MHWS and EHWS. (b) Mangroves on the high tidal flats usually between MSL and MHWS, bordered in this example by
saltmarsh on the landward side. (c) Seagrass between MLWN and subtidal zone, and saltmarsh between MHWS and EHWS.

Tidal Flat, Figure 3 Idealized diagram showing a range of tidal flats underlain by mud, or mud grading downslope to sand flats,
muddy tidal flat grading upslope to sand flats fronting a low-gradient beach or sandy spit/chenier, and sandy tidal flat fronting
a low-gradient beach or sandy spit/chenier.

located in seaward parts of an estuary tend to be more saline
in groundwater and pellicular water than those located in
the riverine parts; (2) location on the tidal flat relative to
MSL; (3) the stage of the tidal cycle in terms of spring tide
versus neap tide, and rising tidal water versus falling tidal
water; (4) the length of exposure of the tidal flat within
the tidal cycle or between spring tides and neap tide phases;
and (5) the climate, which determines the extent of evapo-
ration, the influence of rainfall on the tidal flat, and the
development of mangroves and saltmarsh. The salinity of
the tidal flat groundwater tends to be relatively constant
from tide to tide during a given season and can even be con-
stant over the entire year. Transpiration by tidal flat vegeta-
tion increases baseline salinities of pellicular and
groundwaters.
While there may be a general trend of increasing tidal
flat salinity from low tidal levels to the high tide mark, this
can be reversed by freshwater seepage along the margin of
668
Tidal Flat, Figure 4 Diagram showing the typical geomorphic and sedimentologic location of tidal flats within an idealized estuary.
the hinterland, from beaches, and from spits and cheniers
(Semeniuk, 1983). Freshwater seepage can manifest as
discharge along the upper tidal flat surface as surface
flows, springs, or as subterranean flow along appropriate
stratigraphic conduits or sheets (Semeniuk, 1983;
Cresswell, 2000; Semeniuk et al., 2011).
Tidal flats in an estuarine environment
Tidal flats occur in different and varied geomorphic and
sedimentologic settings within an estuary. The main
TIDAL FLAT
settings are platforms bordering the margins of estuaries,
tidal parts of flood-tidal deltas, tidal parts of intra-
estuarine riverine deltas, the crests and flanks of middle-
of-estuary shoals, and the protected lagoons leeward of
spits. Figure 4 diagrammatically shows the locations of
tidal flats within an idealized estuary.
The platforms bordering the margins of estuaries, since
they can occur continuously or discontinuously along the
whole length of an estuary, can traverse salinity fields from
marine (at seaward parts of an estuary) to brackish or even
freshwater (at riverine parts of an estuary). In terms of
 TIDAL FLAT
substrates, they can be sandy, muddy sand, or muddy,
depending on sediment supply and local hydrodynamics.
The tidal parts of flood-tidal deltas tend to be underlain
by sand and have marine salinities. The tidal parts of
intra-estuarine riverine deltas can be brackish to freshwater,
varying between the seasons (and even marine water during
the dry season) and, depending on the nature of the delta,
can be underlain by sand, muddy sand, or mud (see Estua-
rine Deltaic Wetlands and Deltas). The crests and flanks of
middle-of-estuary shoals reside in water that varies from
marine at seaward parts of the estuary to brackish/freshwa-
ter in riverine parts of an estuary. Again, depending on sed-
iment supply, the tidal flats can be underlain by sand,
muddy sand, or mud. The tidal parts of protected lagoons,
leeward of spits, tend to be in seaward parts of the estuary
(though they can occur widely) and, as such, waters usually
are marine salinity. Their substrates are sandy. However, if
there is abundant mud in suspension in the estuarine waters,
the low-energy protected leeward zone results in mud accu-
mulation and the tidal flats are mud floored.
With all these tidal flats, except generally for the flood-
tidal delta, if there is both sand and mud available, the sand
and mud commonly are partitioned in different parts of the
environment because of hydrodynamic processes, and as
such, there can be depth-related mud tidal flats grading
to sand tidal flats.
Biota of tidal flats within estuaries
There are various factors that influence the structure and
composition of biota that inhabit a tidal flat. These include
biogeography and climate, inundation frequency, sub-
strates, moisture, salinity, hydrochemistry, herbivory, and
predation (Verwey, 1954; Paine, 1974; Lubchenco, 1978;
Semeniuk, 1983; Reise, 1985; Reise et al., 1994; Reise,
2000; Cronin et al., 2010).
Biogeography and climate are major determinants of
the composition of biota of tidal flats in estuaries. The
range of biota includes mangroves and their associated
fauna in tropical and subtropical environments, rushes,
sedges, samphire, and other saltmarsh plants and their
associated fauna in tropical, subtropical, and temperate
environments, with different species in the various bio-
geographic regions (Tomlinson, 1986; Adams, 1990;
Pen et al., 2000; Cresswell and Semeniuk, 2011), and var-
ious crustacean-polychaete-mollusc assemblages in tropi-
cal, subtropical, and temperate environments, again, with
different species in the various biogeographic regions
(Wolff, 1983; White, 1989; Semeniuk et al., 2000;
Brearley, 2005). For instance, western hemisphere man-
groves have different species than eastern hemisphere
mangroves (Tomlinson, 1986), and their composition,
structure, and physiognomy change with climate and lati-
tude (Tomlinson, 1986; Cresswell and Semeniuk, 2011;
Semeniuk and Cresswell, 2013).
Estuarine tidal flat biota comprise plants, resident
fauna, invading and temporary fauna, and microbiota.
The plants of tidal flats can be categorized into four main
669
groups, viz., mangroves, saltmarsh, seagrass, and algae.
Mangroves occupy the interval between MSL and MHWS
but, in more humid and tropical climates, occupy the inter-
val between MSL and EHWS. Saltmarsh occupies the
interval between MSL and EHWS, but depending on cli-
mate, freshwater seepage, and tidal range, saltmarsh can
more narrowly occupy the interval between MHWS and
EHWS. Seagrass and algae generally occur in lower parts
of the tidal flat and extend to subtidal zones.
Within a given biogeographic region, the composition
of tidal flat biota, in site-specific locations, is related to
tidal zones (inundation frequency), substrate type, sub-
strate moisture, and salinity, commonly resulting in spe-
cies zonation (Bridgewater, 1975; Wolff, 1983;
Semeniuk, 1985; White, 1989; Pennings and Callaway,
1992; Emery et al., 2001; Pennings et al., 2005; Silvestri
et al., 2005; and Semeniuk and Cresswell, this volume,
see Species Zonation). Mangrove and saltmarsh emergent
vegetation conspicuously exhibit species zonation
(Semeniuk et al., 1978; Tomlinson, 1986; Adam, 1990;
Cresswell and Semeniuk 2011).
There is a specialized and often unique fauna associated
with mangroves, saltmarsh, and seagrass. For instance,
in mangroves, there is a biogeography-specific and
climate-specific mangrove-associated infauna of crusta-
cea and polychaetes; an epifauna of crustacea (crabs, bar-
nacles), insects, and vertebrates; and a canopy fauna of
avifauna, reptiles, insects, and mammals (bats) (MacNae,
1968; Semeniuk et al., 1978; Hutchins and Saenger,
1987; Johnstone, 1990; Kathiresan and Bingham, 2001).
Similarly in saltmarshes, there is a biogeography-specific
and climate-specific saltmarsh-associated fauna, but also
a partitioning of fauna relative to position of tidal levels
and salinity. Adam (1990) uses several approaches to cat-
egorizing saltmarsh fauna: taxonomically, ecologically
(e.g., marine, freshwater, and terrestrial affinities), trophi-
cally, subhabitat occupied, and residence status. For
instance, for a given biogeographic and climate setting,
saltmarsh may have an infauna of crustacea, other arthro-
pods, molluscs, polychaetes, and meiofauna; an epifauna
of crustacea, other arthropods, molluscs, insects, and ver-
tebrates; and a foliage-associated fauna of avifauna, rep-
tiles, mammals, gastropods, and insects (Adam, 1990).
The resident fauna of tidal flats outside of the zone of
mangroves and saltmarsh, as macrofauna, can be diverse
and complex in composition and its internal dynamics.
Examples of compositional variability and complex
dynamics are provided by Beukema (1976, 1981),
Virnstein (1977), Bell and Coull (1978), Quammen
(1982), Reise (1982, 1991, 2000), Ambrose (1984), Bot-
tom (1984), Tsuchiya and Nishihira (1986), Dittmann
(1990, 1995, 1996, 2000, 2002), Nehls and Thiel (1993),
Jones et al. (1994), Thiel and Dernedde (1994), Vargas
(1988), Ragnarsson and Raffaelli (1999), Attrill
et al. (2001), Dittmann and Vargas (2001), Hagberg et al.
(2004), Reichert and Buchholz (2006), Büttger
et al. (2008), and Buschbaum et al. (2009). In the first
instance, such fauna can be categorized as epifauna
670 TIDAL FLAT
Tidal Flat, Figure 5 Diagram showing blocks of sediment with different types of biota commensurate with substrate type and tidal
level setting.
(inhabiting the tidal flat surface; this includes grazing gas-
tropods, scavenging echinoderms, sessile mussels,
encrusting oysters), infauna (inhabiting the tidal flat sub-
surface in structured burrows, or cavities, or simply bur-
ied; this includes bivalves, polychaetes, echinoderms,
and crustacea; endobenthic is a term also applied to
infauna), or alternating infauna/epifauna (inhabiting the
subsurface but emerging periodically onto the surface for
feeding or breeding; this includes crabs, goby fish, some
polychaetes, and some gastropods, e.g., Unno and
Semeniuk, 2011). Again, these assemblages are biogeog-
raphy specific and climate specific in composition and
structure but also substrate specific in that low tidal sand
flat fauna can be markedly different from low tidal mud
flat fauna for a given biogeographic and climate setting.
A summary diagram of the complex associations of biota
on unvegetated low tidal flats, in mangrove zones, and in
saltmarsh is presented in Figure 5. A summary of the dis-
tribution, zoning, and environment-specific burrow struc-
tures (ichno-structures) developed across tidal flats by
various fauna, and their development of structures is pro-
vided by Reineck and Singh (1980).
Microbiota of tidal flats includes diatoms, foraminifera,
amoeba, bacteria, fungi, and small-scaled invertebrates
such as annelids, nematodes, turbellarians, ostracods, crus-
tacean larvae, insect larvae, and juvenile molluscs, among
others (Higgins and Thiel, 1988). The small-sized fauna
that lives interstitial to sand in sandy habitats and that gen-
erally passes through a 1 mm mesh (but retained by
a 30–45 mm mesh) in sandy and muddy habitats is termed
meiofauna. Meiofauna and other microbiota are abundant
in tidal flat sediments as epibiota and endobiota, often
zonally distributed, and may be vertically mobile with the
rise and fall of the tide and groundwater under tidal
flats (Wood, 1967; Happey-Wood and Jones, 1988;
Brotas and Plante-Cuny, 1998; Hortona et al., 1999;
Haslett, 2001; Mitbavkar and Anil, 2002, 2004; Coull,
2009; Ghosh, 2012; Hankin et al. (2012). They are com-
monly the food source for grazing and sediment-foraging
invertebrates.
A special resident fauna on tidal flats includes some
species of molluscs and worms (mussels, oysters, serpulid
worms) that form dense colonies resulting in skeletal
structures that cover the tidal flats as reefal biostromes or
locally emerge as reefal bioherms (skeletal buildup terms
from Nelson et al., 1962). Oysters and mussels typically
can form sheetlike skeletal structures (or reefal
biostromes) that cover the tidal flat in patches or ribbons
 TIDAL FLAT
(Kuenen, 1942; van Straaten and Kuenen, 1957; Bosence,
1979; Dittmann, 1990; Nehls and Thiel, 1993; Eisma,
1998). Where hard surfaces are developed (such as mussel
beds, oyster reefs, serpulid reefs, shell-lag pavements, or
scattered shells on the sediment surface), encrusting
organisms and (micro-scaled) hard-surface communities
such as oysters, hydrozoans, anthozoans, bryozoans, and
algae can develop (Albrecht, 1998).
The invading or temporary fauna of tidal flats, i.e., fauna
that is not permanently resident on the tidal flat, include fish
(including rays), crabs, reptiles, and octopus that invade the
tidal flat on a high tide to feed; birds, reptiles, and mammals
that invade the tidal flat on a low tide to feed; and the birds
that use the saltmarsh vegetation and mangrove canopy for
nesting, feeding, roosting, and loafing.
Figure 6 illustrates in generalized form the contrast
between biota across a tropical macrotidal sandy to muddy
tidal flat and that of a tropical macrotidal wholly muddy
tidal flat, and the contrast between biota in various
temperate-climate mesotidal and microtidal sandy tidal
flats. Figure 7 illustrates various habitats and ecological/
biological aspects of tidal flats. Figure 8 shows some of
the biological aspects of the tidal flats.
Tidal flats as ecological systems – complexity, and
trophic and ecologic function
Tidal flats are complex systems – from the large scale of
geomorphology, habitats, ecosystems, geochemistry,
hydrology, and hydrochemistry to the smaller scales of
autoecology and microbial processes. Occurring in the
coastal zone, as the low-gradient interface between ocean,
land, atmosphere, and groundwater, they support
a diversity of biota and a diversity of physical, chemical,
and biological processes and, as such, function as
a storage system for living and decaying biomass and for
biochemical/geochemical products. As a coastal system
bridging the environments of ocean and land, and in estu-
arine settings bridging the environments of freshwater and
marine water, tidal flats also provide a multiplicity of eco-
logical functions. As a major zone of primary and second-
ary productivity, they are an important food source for
marine and terrestrial animals.
In the first instance, there is primary production by ben-
thic plant life such as mangroves, saltmarsh vegetation,
seagrasses, algae, and diatoms and, during the high tide,
primary production in the water column by phytoplankton.
Plant biomass is then cycled through the tidal ecosystem by
herbivory and/or nectivory by insects, gastropods, crabs,
turtles, bats, fish, and mammals, leaf litter fall and death
and decay of plant matter, and filter feeding by bivalves
and other invertebrates. Plant material in decayed or com-
minuted form accumulates as detritus where it is either
ingested by detritus feeders (gastropods, polychaetes, crus-
tacea) or transformed by microbiota to microbial slime and
grazed by gastropods, polychaetes, and crustacea, or
(if mobilized into the water column) extracted by filter
feeders, or where it is incorporated on and into the
671
sediments becomes involved in complex geochemical and
biochemical transformations and recycled by microbiota
as part of the nitrogen, sulfur, phosphorus, and carbon cycle
(see “Mineralization”) and transformed into inorganic
products. Following incorporation of biomass and detritus
(in various stages of decay) into herbivores, nectivores,
and filter feeders, these primary consumers are preyed on
by predators such as gastropods, echinoderms, nektonic
fish, demersal fish, stingrays, octopus, and terrestrial
invaders such as snakes, other reptiles, mammals, and wad-
ing birds.
As such, tidal flats are major feeding grounds for nek-
ton and demersal fish on the high tide and major feeding
grounds for reptiles, mammals, and avifauna on the low
tide. Figure 9 diagrammatically shows the ecological
function of mangroves on a tidal flat, with blocks of sedi-
ment showing different types of biota and their activity
commensurate with substrate type and tidal level setting,
and the invasion of terrestrial animals on the low tide
and marine animals on the high tide to feed on the resident
biota of the tidal flat.
Examples specifically of bird life utilizing the tidal flats,
their dependence on the ecosystem and effects on the eco-
system, and their nesting in mangroves and saltmarsh are
provided by Semeniuk et al. (1978), Goss-Custard (1980),
Zwarts (1981), Smit (1984), Johnstone (1990), Dann
(1991), Beukema et al. (1993, 2010), Nehls and Tiedemann
(1993), Piersma et al. (1993), Ens et al. (1994), Goss-
Custard et al. (1996), Backwell et al. (1998), Raines
et al. (2000), Connolly and Colwell (2005), Granadeiro
et al. (2007), and Spalding et al. (2010).
Tidal flats and estuarine shallow waters also are consid-
ered to be nurseries and sanctuaries for fauna (McHugh,
1967; Blaber and Blaber, 1980; Staples, 1980; Haedrich,
1983; Boehlert and Mundy, 1988; Beukema, 1992; Maes
et al., 1998; Morrison et al., 2002; Francis et al., 2005;
Reise et al., 2010) – for contrasting discussions of the
importance of shallow-water habitats for fish nurseries,
see Able (2005), Baker and Sheaves (2005, 2007), and
Johnston et al. (2007). While shallow waters are consid-
ered to be nurseries for fish and juvenile crabs and
shrimps, nurseries also are specifically provided on tidal
flats themselves by pools of water that are left stranded
on the low tide. During periods of recruitment, the numer-
ous small water pools on uneven surfaces of open tidal
flats, in mangroves, and in saltmarsh are abundant with
small fish. Sanctuaries are provided in the mangroves,
saltmarsh, and seagrass.
Tidal flats provide variable habitats for organisms
related to sediment types, tidal levels, and hydrochemistry
and, as such, for a given biogeography and climate setting,
provide the basis for biodiversity.
Summary
Tidal flats are tidally exposed surfaces, underlain by
a variety of substrates, and variably vegetated depending
on climate and tidal level. The entire surface between
672 TIDAL FLAT

Tidal Flat, Figure 6 Profiles of some typical estuarine tidal flats showing substrate types and generalized and simplified composition
of biota (mangrove, saltmarsh, invertebrates, mussels). Information on fauna in Wadden Sea from Wolff (1983); in the Lawley River
Estuary and King Sound from Semeniuk (1981), Wells and Slack-Smith (1981), and Semeniuk (unpublished data); and in Leschenault
Inlet Estuary from Semeniuk and Wurm (2000), Semeniuk (2000), and Dürr and Semeniuk (2000).
 TIDAL FLAT 673

Tidal Flat, Figure 7 Various tidal flats in estuaries and some of their ecological/biological aspects. (a) Sandy low tidal flat, with
low-relief shore-parallel shoals and depressions (small-scale habitats), in a northern Kimberley estuary, northwestern Australia.
(b) Zoned mangroves on a muddy tidal flat in the northern Cambridge Gulf estuary, northwestern Australia; landward of the
mangroves is a vegetation-free saline muddy flat (salt flat); spit emanates from the rocky spur and crosses the salt flat. (c) Zoned
mangroves on a muddy tidal flat in northern King Sound, northwestern Australia; seaward of the mangroves is a moderately sloping
mud flat, and landward is a vegetation-free muddy salt flat; also on the salt flat are sand bodies (cheniers). (d) Saltmarsh along shore
of the Leschenault Inlet Estuary, southwestern Australia, showing three vegetation zones – shoreline rush in immediate foreground,
samphire in foreground to middle ground (alternating bands of pink, green, and grey), and samphire in the background (pink);
Avicennia marina shrub occurs in seaward saltmarsh zone. (e) Low tidal muddy sand flat adjoining a sloping sandy and shelly beach,
Tasmania, southeastern Australia. (f) Low tidal muddy flat adjoining a mud-floored mangrove fringe, Botany Bay, southeastern
Australia; the mud flat is pocked with worm burrows, crab burrows, and fish-feeding excavations. (g) Slightly muddy sand low tidal
flat, Tasmania, southeastern Australia; soldier crabs swarming and feeding on surface. (h) Low tidal sandy flat with soldier crab
feeding pellets on the surface, Tasmania, southeastern Australia; in background is a shore-parallel low-relief shoal; in foreground, the
shiny surface is due to a thin film of mud giving the impression the tidal flat is a mud flat.

ELWS and EHWS is the tidal zone, and any low-gradient
surface between these levels is a tidal flat. Tidal flats are
not always simple uniformly sloping surfaces; they often
have variation as determined by hydrodynamic setting,
e.g., low hummocks of sand on mud flats, low relief
cheniers, or tidal creeks, as well as variable substrates.
These smaller-scale variations influence the development
of small-scale habitats and the distribution of biota across
the tidal flat. In providing variable habitats for organisms
related to sediment types, tidal levels, and hydrochemistry,
for a given biogeography and climate setting, tidal flats
form the foundation for coastal biodiversity.
Tidal flats occur in varied geomorphic and sedimento-
logic settings within an estuary; these include platforms
bordering estuary margins, tidal parts of flood-tidal deltas,
tidal parts of intra-estuarine riverine deltas, the crests and
flanks of middle-of-estuary shoals, and the protected
lagoons leeward of spits. Where sand and mud are avail-
able, the sediments commonly are partitioned across the
tidal flat because of hydrodynamic processes, resulting
in depth-related mud flat grading to sand flats.
When exposed at low tide, tidal flats are subject to
evaporation and transpiration, and a salinity gradient from
seaward to landward is developed due to the interacting
674 TIDAL FLAT

Tidal Flat, Figure 8 Biology of some tidal flats of estuaries in southeastern Australia. (a) Polychaete workings of sediment (low-relief
conical mounds) and burrows on a low tidal flat, Tasmania. (b) Fish and stingray feeding excavations on a rippled low tidal sandy flat,
Tasmania. (c) Slightly muddy sand low tidal flat with shallow pools, Tasmania; swarm of soldier crabs on the surface. (d) Swarm of
soldier crabs on low tidal flat and white-faced heron (Egretta novaehollandiae) feeding on the crabs, Botany Bay, southeastern
Australia.

Tidal Flat, Figure 9 Diagram showing ecological function of mangroves on a tidal flat with blocks of sediment showing different
types of biota commensurate with substrate type and tidal level setting (modified after Semeniuk et al., 1978).
 TIDAL FLAT
effects of inundation, evaporation, transpiration, and sedi-
ment types. The increasing salinity can be locally diluted
by freshwater seepage from the margin of the hinterland,
beaches, and spits and cheniers.
Estuarine tidal flat biota comprise plants, resident fauna,
invading and temporary fauna, and microbiota. At the over-
arching scale, biogeography and climate are major determi-
nants of the composition of tidal flat biota. Tidal flat biota
includes mangroves and their associated fauna in tropical
and subtropical climates, and rushes, sedges, samphire,
and other saltmarsh plants and their associated fauna in
tropical, subtropical, and temperate climates. Mangrove
and saltmarsh emergent vegetation conspicuously exhibits
species zonation. A range of mollusc, crustacean, and poly-
chaete fauna occurs in tidal flat habitats, the composition of
which is related to tidal zones, substrate type, substrate
moisture, and salinity, and commonly results in species
zonation. The resident fauna of tidal flats can be diverse
and complex in composition and its internal population
dynamics and ecological dynamics. It can be categorized
as epifauna, infauna, or alternating infauna/epifauna and
is biogeography specific, climate specific, and substrate
specific in composition and structure. The invading or tem-
porary fauna of tidal flats are variable in terms of taxonomic
group; they include fish (including rays), crabs, reptiles,
and octopus that invade tidal flats on a high tide to feed;
birds, reptiles, and mammals that invade tidal flats on
a low tide to feed; and the birds that use the saltmarsh veg-
etation and mangrove canopy for a variety of activities.
A special resident fauna on tidal flats of mussels, or
oysters, or serpulid worms can form dense colonies
resulting in skeletal structures that cover tidal flats as
reefal biostromes or locally emerge as reefal bioherms.
Where hard surfaces are developed, encrusting organisms
and (micro-scaled) hard-surface communities of oysters,
hydrozoans, anthozoans, bryozoans, and algae can
develop.
Microbiota of tidal flats includes diatoms, foraminifera,
amoebae, bacteria, fungi, and small-scaled invertebrates
that are abundant in tidal flat sediments, often zonally dis-
tributed, and vertically mobile with the fluctuation of the
tide and groundwater of tidal flats. They are commonly
the food source for grazing and sediment foraging
invertebrates.
Ecologically, tidal flats are complex systems, even if
substrates appear relatively uniform (e.g., wholly muddy
tidal flats or wholly sandy tidal flats). Complexity is
underpinned by inundation gradients and salinity which
will determine zonation and hence microecology pro-
cesses. Where substrates are graded, or where there is var-
iable low-relief geomorphic expression, and where
freshwater is discharging into the tidal flat, the complexity
of tidal flats increases. Occurring in the coastal zone, as a -
low-gradient interface between ocean, land, atmosphere,
and groundwater, they support a diversity of biota and
a diversity of physical, chemical, and biological processes,
and they function as a storage system for living and
decaying biomass and for biochemical/geochemical
675
products. As coastal systems bridging the environments
of ocean and land and in estuarine settings bridging fresh-
water and marine water, tidal flats also provide
a multiplicity of ecological functions. As major zones of
primary and secondary productivity, they are important
food sources for marine and terrestrial animals.
On tidal flats, there is primary production carried out by
benthic plant life such as mangroves, saltmarsh vegetation,
seagrasses, algae, and diatoms and, during the high tide in
the water column, by phytoplankton. Plant biomass is
cycled though the ecosystem by herbivory and/or nectivory,
leaf litter fall, decay of plant matter, and filter feeding by
invertebrates. Following incorporation of biomass and
detritus by primary consumers, predators carry the biomass
into higher trophic levels. Tidal flats are major feeding
grounds for nekton and demersal fish on the high tide and
for reptiles, mammals, and avifauna on the low tide. Tidal
flats, and in particular mangroves, saltmarsh, and seagrass,
also provide nurseries and sanctuaries for fauna.
Bibliography
Able, K. W., 2005. A re-examination of fish estuarine dependence:
evidence for connectivity between estuarine and ocean habitats.
Estuarine, Coastal and Shelf Science, 64, 5–17.
Adam, P., 1990. Saltmarsh Ecology. Cambridge: Cambridge Uni-
versity Press.
Albrecht, A. S., 1998. Soft bottom versus hard rock: community
ecology of macro-algae on intertidal mussel beds in the Wadden
Sea. Journal of Experimental Marine Biology and Ecology, 229,
85–109.
Ambrose, W. G., 1984. Role of predatory infauna in structuring
marine soft-bottom communities. Marine Ecology Progress
Series, 17(2), 109–115.
Attrill, M. J., Stafford, R., and Rowden, A. A., 2001. Latitudinal
diversity patterns in estuarine tidal flats: indications of global
cline. Ecography, 24, 318–324.
Backwell, P. R. Y., O’Hara, P. D., and Christy, J. H., 1998. Prey
availability and selective foraging in shorebirds. Animal Behav-
iour, 55, 1659–1667.
Baker, R., and Sheaves, M., 2005. Redefining the piscivore assem-
blage of shallow estuarine nursery habitats. Marine Ecology Pro-
gress Series, 291, 197–213.
Baker, R., and Sheaves, M., 2007. Shallow-water refuge paradigm:
conflicting evidence from tethering experiments in a tropical
estuary. Marine Ecology Progress Series, 349, 13–22.
Bell, S., and Coull, B., 1978. Field evidence that shrimp predation
regulates meiofauna. Oecologia, 35, 141–148.
Beukema, J. J., 1976. Biomass and species richness of the macro-
benthic animals living on the tidal flats of the Dutch Wadden
Sea. Netherlands Journal of Sea Research, 10, 236–261.
Beukema, J. J., 1981. The role of the larger invertebrates in the
Wadden Sea ecosystem. In Dankers, N., et al. (eds.), Inverte-
brates of the Wadden Sea. Rotterdam: A. A. Balkema,
pp. 211–221.
Beukema, J. J., 1992. Dynamics of juvenile shrimp Crangon
crangon in a tidal-flat nursery of the Wadden Sea after
mild and cold winters. Marine Ecology Progress Series, 83,
157–165.
Beukema, J. J., Essink, K., Michaelis, H., and Zwarts, L., 1993.
Year-to-year variability in the biomass of macrobenthic animals
on tidal flats of the Wadden Sea: How predictable is this food
source for birds? Journal of Sea Research, 31, 319–330.
676 TIDAL FLAT
Blaber, S. J. M., and Blaber, T. G., 1980. Factors affecting the distri-
bution of juvenile estuarine and inshore fish. Journal of Fish
Biology, 17, 143–162.
Boehlert, G. W., and Mundy, B. C., 1988. Roles of behavioral and
physical factors in larval and juvenile fish recruitment to estua-
rine nursery areas. In Weinstein, M. P. (ed.), Larval Fish and
Shellfish Transport Through Inlets. Bethesda: American Fisher-
ies Society. American Fisheries Society Symposium, Vol.
3, pp. 51–67.
Bosence, D. W. J., 1979. The factors leading to aggregation and reef
formation in Serpula vermicularis L. In Larwood, G., and Rosen,
B. R. (eds.), Biology and Systematics of Colonial Organisms.
London: Academic.
Bottom, M. L., 1984. The importance of predation by horseshoe
crabs, Limulus polyphemus, to an intertidal sand flat community.
Journal of Marine Research, 42, 139–161.
Brearley, A., 2005. Ernest Hodgkin’s Swanland: Estuaries and
Coastal Lagoons of South-western Australia. Crawley: Univer-
sity of Western Australia Press.
Bridgewater, P. B., 1975. Peripheral vegetation of Westernport Bay.
Proceedings of the Royal Society of Victoria, 87, 69–78.
Brotas, V., and Plante-Cuny, M. R., 1998. Spatial and temporal pat-
terns of microphytobenthic taxa of estuarine tidal flats in the
Tagus Estuary (Portugal) using pigment analysis by HPLC.
Marine Ecology Progress Series, 171, 43–57.
Buschbaum, C., Dittmann, S., Hong, J.-S., Hwang, I.-S., Strasser,
M., Thiel, M., Valdivia, N., Yoon, S.-P., and Reise, R., 2009.
Mytilid mussels: global habitat engineers in coastal sediments.
Helgolanderand Marine Research, 63, 47–58, doi:10.1007/
s10152-008-0139-2.
Büttger, H., Asmus, H., Asmus, R., Buschbaum, C., Dittmann, S.,
and Nehls, G., 2008. Community dynamics of intertidal soft-
bottom mussel beds over two decades. Helgolanderand Marine
Research, 62, 23–36.
Cameron, W. M., and Pritchard, D. W., 1963. Estuaries. In Hill,
M. N. (ed.), The Sea. New York: Wiley, Vol. 2, pp. 306–324.
Cartwright, D. E., 1999. Tides – A Scientific History. Cambridge:
Cambridge University Press.
Connolly, L. M., and Colwell, M. A., 2005. Comparative use of
longline oysterbeds and adjacent tidal flats by waterbirds. Bird
Conservation International, 15, 237–255.
Coull, B. C., 2009. Role of meiofauna in estuarine soft-bottom hab-
itats. Austral Ecology, 24(4), 327–343.
Cresswell, I. D., 2000. Ecological significance of freshwater seeps
along the western shore of the Leschenault Inlet estuary. Journal
of the Royal Society of Western Australia Special Issue on the
Leschenault Inlet Estuary, 83, 285–292.
Cresswell, I. D., and Semeniuk, V., 2011. Mangroves of the Kimber-
ley Coast, Western Australia: ecological patterns in a tropical ria
coast setting. Journal of the Royal Society of Western Australia,
94, 213–237.
Cronin, J. P., Tonsor, S. J., and Carson, W. P., 2010. A simultaneous
test of trophic interaction models: which vegetation characteris-
tic explains herbivore control over plant community mass? Ecol-
ogy Letters, 13, 202–212.
Dann, P., 1991. Feeding behaviour and diet of the Double-banded
Plovers Charadrius bicinctus in Westernport, Victoria. Emu,
91, 179–184.
Dittmann, S., 1990. Mussel beds – amensalism or ameleorisation
for intertidal fauna? Helgolander Meeresunters, 44, 335–352.
Dittmann, S., 1995. Benthos structure on tropical tidal flats of Aus-
tralia. Helgoländer Meeresunters, 49, 539–551.
Dittmann, S., 1996. Effects of macrobenthic burrows on infaunal
communities in tropical tidal flats. Marine Ecology Progress
Series, 134, 119–130.
Dittmann, S., 2000. Zonation of benthic communities in a tropical tidal
flat of north-east Australia. Journal of Sea Research, 43, 33–51.
Dittmann, S., 2002. Benthic fauna in tropical tidal flats –
a comparative perspective. Wetlands Ecology & Management,
10, 189–195.
Dittmann, S., and Vargas, J. A., 2001. Tropical tidal flat benthos
compared between Australia and Central America. In Reise,
K. (ed.), Ecological Comparisons of Sedimentary Shores. Ber-
lin: Springer. Ecological Studies, Vol. 151, pp. 275–293.
Dürr, V., and Semeniuk, T. A., 2000. Long-term spatial dynamics of
polychaetes in Leschenault Inlet estuary. Journal of the Royal
Society of Western Australia Special Issue on the Leschenault
Inlet Estuary, 83, 429–441.
Eisma, D., 1998. Intertidal Deposits: River Mouths, Tidal Flats, and
Coastal Lagoons. Boca Raton: CRC Press.
Emery, N. C., Ewanchuk, P. J., and Bertness, M. D., 2001. Compe-
tition and salt-marsh plant zonation: stress tolerators may be
dominant competitors. Ecology, 82, 2471–2485.
Ens, B. J., Piersma, T., and Drent, R. H., 1994. The dependence of
waders and waterfowl migrating along the East Atlantic Flyway
on their coastal food supplies: what is the most profitable
research program? Ophelia: International Journal of Marine
Biology, Supplement, 6, 127–151.
Francis, M. P., Morrison, M. A., Leathwick, J., Walsh, C., and Mid-
dleton, C., 2005. Predictive models of small fish presence and
abundance in northern New Zealand harbours. Estuarine,
Coastal and Shelf Science, 64, 419–435.
Ghosh, A., 2012. Estuarine foraminifera from the Gulf of Cambay.
Journal Geological Society of India, 80, 65–74.
Goss-Custard, J. D., 1980. Competition for food and interference
among waders. Ardea, 68, 31–52.
Goss-Custard, J. D., McGrorty, S., and dit Durell, S. E. A. le V.,
1996. The effect of Oystercatchers Haematopus ostralegus on
shellfish populations. Ardea, 84A, 453–468.
Granadeiro, J. P., Santos, C. D., Dias, M. P., and Palmeirim, J. M.,
2007. Environmental factors drive habitat partitioning in birds
feeding in intertidal flats: implications for conservation.
Hydrobiologia, 587, 291–302.
Haedrich, R. L., 1983. Estuarine fishes. In Ketchum, B. H. (ed.),
Estuaries and Enclosed Seas. Ecosystems of the World. Amster-
dam: Elsevier Scientific, Vol. 26, pp. 183–207.
Hagberg, J., Tunberg, B. J., Wieking, G., Kroncke, I., and Belgrano,
A., 2004. Effects of climate variability on benthic communities.
In Stenseth, N. C., and Ottersen, G. (eds.), Marine Ecosystems
and Climate Variation – The North Atlantic – A Comparative
Perspective. New York: Oxford University Press, pp. 115–121.
Hankin, S. L., Weilhoefer, C. L., Kaldy, J. E., and DeWitt, D. H.,
2012. Sediment diatom species and community response to
nitrogen addition in Oregon (USA) estuarine tidal wetlands. Wet-
lands, 32(6), 1023–1031.
Happey-Wood, C. M., and Jones, P., 1988. Rhythms of vertical
migration and motility in intertidal benthic diatoms with particu-
lar reference to Pleurosigma angulatum. Diatom Research, 3,
83–93.
Haslett, S. K., 2001. The palaeoenvironmental implications of the
distribution of intertidal foraminifera in a tropical Australian
estuary: a reconnaissance study. Australian Geographical Stud-
ies, 39, 67–74.
Higgins, R. P., and Thiel, H., 1988. Introduction to the study of
meiofauna. Washington, DC: Smithsonian Institution Press.
Hortona, B. P., Edward, R. J., and Lloyd, J. M., 1999. UK intertidal
foraminiferal distributions: implications for sea-level studies.
Marine Micropaleontology, 36, 205–223.
Hutchins, P., and Saenger, P., 1987. Ecology of Mangroves.
St. Lucia: University of Queensland Press.
 TIDAL FLAT
Johnston, R., Sheaves, M., and Molony, B., 2007. Are distributions
of fishes in tropical estuaries influenced by turbidity over small
spatial scales? Journal of Fish Biology, 71, 657–671.
Johnstone, R. E., 1990. Mangroves and Mangrove Birds. Records of
the Western Australian Museum, Supplement 32. Perth: Western
Australian Museum.
Jones, C. G., Lawton, J. H., and Shachak, M., 1994. Organisms as
ecosystem engineers. Oikos, 69, 373–386.
Kathiresan, K., and Bingham, B. L., 2001. Biology of mangroves
and mangrove ecosystems. Advances in Marine Biology, 40,
81–251.
Kuenen, D. J., 1942. On the distribution of mussels on the intertidal
sand flats near Den Helder. Archives Néerlandaises de Zoologie,
6, 117–160.
Lubchenco, J., 1978. Plant species diversity in a marine intertidal
community: importance of herbivore food preferences and algal
competitive abilities. American Naturalist, 112, 23–29.
Macnae, W., 1968. A general account of the fauna and flora of man-
grove swamps and forests in the Indo-West-Pacific region.
Advances in Marine Biology, 6, 73–270.
Maes, J., Taillieu, A., van Damme, P. A., Cottenie, K., and Ollevier,
F., 1998. Seasonal patterns in the fish and crustacean community
of a turbid temperate estuary (Zeeschelde Estuary, Belgium).
Estuarine, Coastal and Shelf Science, 47, 143–151.
McHugh, J. L., 1967. Estuarine nekton. In American Association
for the Advancement of Science. Washington: Publication 83
(Estuaries), pp. 581–620.
Mitbavkar, S., and Anil, A. C., 2002. Diatoms of the microphy-
tobenthic community: population structure in a tropical intertidal
sand flat. Marine Biology, 140, 41–57.
Mitbavkar, S., and Anil, A. C., 2004. Vertical migratory rhythms of
benthic diatoms in a tropical intertidal sand flat: influence of irra-
diance and tides. Marine Biology, 145, 9–20.
Morrison, M. A., Francis, M. P., Hartill, B. W., and Parkinson,
D. M., 2002. Diurnal and tidal variation in the abundance of
the fish fauna of a temperate tidal mudflat. Estuarine, Coastal
and Shelf Science, 54, 793–807.
Nehls, G., and Thiel, M., 1993. Large-scale distribution patterns
of the mussel Mytilus edulis in the Wadden Sea of Schleswig–
Holstein: do storms structure the ecosystem? Netherlands
Journal of Sea Research, 31, 181–187.
Nehls, G., and Tiedemann, R., 1993. What determines the densities
of feeding birds on tidal flats? A case study on dunlin, Calidris
alpina, in the Wadden Sea. Netherlands Journal of Sea
Research, 31, 375–384.
Nelson, H. F., Brown, C. M. W., and Brineman, J. H., 1962. Skeletal
limestone classification. In Ham, W. E. (ed.), Classification of
Carbonate Rocks: A Symposium. American Association of
Petroleum Geologists Memoir 1, Oklahoma, pp. 62–84.
Paine, R. T., 1974. Intertidal community structure: experimental
studies on the relationship between a dominant competitor and
its principal predator. Oecologia, 15, 93–120.
Pen, L., Semeniuk, V., and Semeniuk, C. A., 2000. Peripheral wet-
land habitats and vegetation of Leschenault Inlet estuary. Jour-
nal of the Royal Society of Western Australia Special Issue on
the Leschenault Inlet Estuary, 83, 293–316.
Pennings, S. C., and Callaway, R. M., 1992. Salt marsh plant zona-
tion: the relative importance of competition and physical factors.
Ecology, 73, 681–690.
Pennings, S. C., Grant, M.-B., and Bertness, M. D., 2005. Plant
zonation in low-latitude salt marshes: disentangling the roles of
flooding, salinity and competition. Journal of Ecology, 93,
159–167.
Piersma, T., de Goeij, R., and Tulp, T., 1993. An evaluation of inter-
tidal feeding habitats from a shorebird perspective: towards
677
relevant comparisons between temperate and tropical mudflats.
Netherlands Journal of Sea Research, 31, 503–512.
Quammen, M. L., 1982. Influence of subtle substrate differences on
feeding by shorebirds on intertidal mudflats. Marine Biology, 71,
339–343.
Ragnarsson, S., and Raffaelli, D. G., 1999. Effects of the mussel
Mytilus edulis on the invertebrate fauna of sediments. Journal
of Experimental Marine Biology & Ecology, 241, 31–44.
Raines, J., Youngson, K., and Unno, J., 2000. Use of the
Leschenault Inlet estuary by waterbirds. Journal of the Royal
Society of Western Australia Special Issue on the Leschenault
Inlet Estuary, 83, 503–512.
Reichert, K., and Buchholz, F., 2006. Changes in the
macrozoobenthos of the intertidal zone at Helgoland (German
Bight, North Sea): a survey of 1984 repeated in 2002.
Helgolander Marine Research, 60, 213–223.
Reineck, H. E., and Singh, I. B., 1980. Depositional Sedimentary
Environments, 2nd edn. Berlin: Springer.
Reise, K., 1982. Long term changes in the macrobenthic inverte-
brate fauna of the Wadden Sea: are polychaetes about to take
over? Netherlands Journal of Sea Research, 16, 29–36.
Reise, K., 1985. Tidal Flat Ecology. Berlin: Springer.
Reise, K., 1991. Macrofauna in mud and sand of tropical and tem-
perate tidal flats. In Elliott, M., and Ducrotoy, J.-P. (eds.), Estuar-
ies and Coasts: Spatial and Temporal Intercomparisons.
Fredensborg: Olsen and Olsen, pp. 211–216.
Reise, K. (ed.), 2000. Ecological Comparisons of Sedimentary
Shores. Berlin: Springer. Ecological Studies, Vol. 151.
Reise, K., Herre, E., and Sturm, M., 1994. Biomass and abundance
of macrofauna in intertidal sediments of Konigshafen in the
northern Wadden Sea. Helgolander Meeresunters, 48, 201–215.
Reise, K., Baptist, M., Burbridge, P., Dankers, N., Fischer, L.,
Flemming, B., Oost, A. P., and Smit, C., 2010. The Wadden
Sea 2010 – A Universally Outstanding Tidal Wetland. Wadden
Sea Ecosystem No. 29. Common Wadden Sea Secretariat
(CWSS), Wilhelmshaven. (ISSN 0946-896X).
Semeniuk, V., 1981. Sedimentology and the stratigraphic sequence
of a tropical tidal flat, North-Western Australia. Sedimentary
Geology, 29, 195–221.
Semeniuk, V., 1983. Mangrove distribution in Northwestern
Australia in relationship to freshwater seepage. Vegetatio, 53,
11–31.
Semeniuk, V., 1985. Development of mangrove habitats along ria
shorelines in north and northwestern Australia. Vegetatio, 60,
3–23.
Semeniuk, T. A., 2000. Small benthic crustacea of the Leschenault
Inlet estuary. Journal of the Royal Society of Western Australia
Special Issue on the Leschenault Inlet Estuary, 83, 429–441.
Semeniuk, V., 2005. Tidal flats. In Schwartz, M. L. (ed.), Encyclo-
paedia of Coastal Science. Berlin: Springer.
Semeniuk, V., and Cresswell, I. D., 2013. A proposed revision of
diversity measures. Diversity, 5, 613–626, doi:10.3390/
d5030613.
Semeniuk, V., and Wurm, P. A. S., 2000. Molluscs of the
Leschenault Inlet estuary – their diversity, distribution, and pop-
ulation dynamics. Journal of the Royal Society of Western Aus-
tralia Special Issue on the Leschenault Inlet Estuary, 83,
377–418.
Semeniuk, V., Kenneally, K. F., and Wilson, P. G., 1978. Mangroves
of Western Australia. Perth: Western Australian Naturalists Club.
Handbook 12.
Semeniuk, V., Semeniuk, T. A., and Unno, J., 2000. The
Leschenault Inlet estuary – an overview. Journal of the Royal
Society of Western Australia Special Issue on the Leschenault
Inlet Estuary, 83, 207–228.
678 TIDAL FLAT SALINITY GRADIENT
Semeniuk, V., Semeniuk, C. A., Tauss, C., Unno, J., and Brocx, M.,
2011. Walpole and Nornalup Inlets: Landforms, Stratigraphy,
Evolution, Hydrology, Water Quality, Biota, and Geoheritage.
Western Australian Museum, Perth (Monograph).
584 p. (ISBN 978-1-920843-37-3).
Silvestri, S., Defina, A., and Marani, M., 2005. Tidal regime, salin-
ity and salt marsh plant zonation. Estuarine Coastal Shelf Sci-
ence, 62, 119–130, doi:10.1016/j.ecss.2004.08.010.
Smit, C. J., 1984. Production of biomass by invertebrates and con-
sumption by birds in the Dutch Wadden Sea areas. In Wolff,
W. J. (ed.), Ecology of the Wadden Sea, 2, part 6. Rotterdam:
Balkema, pp. 290–301.
Spalding, M., Kainuma, M., and Collins, L., 2010. World Atlas of
Mangroves. Malta: Earthscan, Guttenberg Press. ISBN 978-1-
84407-657-4.
Staples, D. J., 1980. Ecology of juvenile and adolescent banana
prawns, Penaeus merguiensis in a mangrove estuary and adja-
cent off-shore area of the Gulf of Carpentaria. Australian Jour-
nal of Marine Freshwater Research, 31, 635–652.
Thiel, M., and Dernedde, T., 1994. Recruitment of shore crabs
Carcinus maenas on tidal flats: mussel clumps as an important
refuge for juveniles. Helgolander Meeresunters, 48, 321–332.
Tomlinson, P. B., 1986. The Botany of Mangroves. Cambridge:
Cambridge University Press.
Tsuchiya, M., and Nishihira, M., 1986. Islands of Mytilus edulis as
a habitat for small intertidal animals: effect of Mytilus age
structure on the species composition of the associated fauna
and community organization. Marine Ecology Progress Series,
31, 171–178.
Unno, J., and Semeniuk, V., 2011. Population dynamics of the
Western Australian soldier crab Mictyris occidentalis Unno
2008 in the Dampier Archipelago - 30 years of observations.
Crustaceana, 84, 905–937.
van Straaten, L. M. J. U., and Kuenen, PH. H., 1957. Accumulation
of fine grained sediments in the Dutch Wadden Sea. Geologie En
Mijnbouw (Nieuwe Series), 19e Jaargang, pp. 329–335.
Vargas, J. A., 1988. Community structure of macrobenthos and the
results of macropredator exclusion on a tropical intertidal mud
flat. Revista Biologia Tropical, 36, 287–308.
Verwey, J., 1954. On the ecology of distribution of cockle and mus-
sel in the Dutch Wadden Sea, their role in sedimentation and the
source of their food supply. Archives Néerlandaises de Zoologie,
10, 171–239.
Virnstein, R. W., 1977. The importance of predation by crabs and fishes
on benthic infauna in Chesapeake Bay. Ecology, 58, 1199–1217.
Wells, F. T., and Slack-Smith, S. M., 1981. Zonation of molluscs in
a mangrove swamp in the Kimberley, Western Australia. West-
ern Australian Museum Publication, pp. 265–274. ISBN
0-7244-8355-1.
White, C. P., 1989. Chesapeake Bay – Nature of the Estuary-
A Field Guide. Centreville: Tidewater Publishers.
Wolff, W. J. (ed.), 1983. Ecology of the Wadden Sea. Rotterdam:
A. A. Balkema.
Wood, E. J. F., 1967. Microbiology of Oceans and Estuaries.
Amsterdam: Elsevier.
Zwarts, L., 1981. Habitat selection and competition in wading birds.
In Smit, C. J., and Wolff, W. J. (eds.), Birds of the Wadden Sea.
Rotterdam: A. A. Balkema, pp. 271–279.
Cross-references
Estuarine Deltaic Wetlands
Mangroves
Saltmarshes
Sandflat
Soldier Crabs (Mictyridae)
Tidal Flat Salinity Gradient
TIDAL FLAT SALINITY GRADIENT
Vic Semeniuk and Christine Semeniuk
V & C Semeniuk Research Group, Warwick, WA,
Australia
Definition
A tidal flat salinity gradient is the systematic or progres-
sive across-tidal-flat change in salinity in groundwater
(as pore water) and pellicular water due to the interacting
effects of inundation, evaporation, transpiration, sediment
types, and freshwater seepage.
Tidal flat salinity
During low tide, when tidal flats are exposed, water under
or within a tidal flat is subject to evaporation and
transpiration, resulting in its salinization, or to dilution
by freshwater seepage. During a low tide, there are two
sub-environments where tidal flat water resides (Figure 1):
1. in a shallow groundwater that is contiguous with the
open waters and whose water table falls and rises with
the rhythm of the tide; at slack-water low tide, this
water table can be
2–10 cm deep on low-gradient
muddy tidal flats and sandy tidal flats or located up to
30 cm depth on more steeply inclined sandy tidal flats;
at extremes, the water table can be at the surface on
very low-gradient tidal flats; the groundwater, which
also can be referred to as the phreatic zone, resides in
the pore spaces of the sediments – this is tidal flat pore
water.
2. as water films circumferential to sediment particles in
the wetted but undersaturated sediment above the
water table during low tide; this can be referred to as
the temporary vadose zone – this is tidal flat pellicular
water.
The salinity of tidal flat groundwater and pellicular
water varies spatially and temporally depending on the
location on the tidal flat relative to mean sea level
(MSL), the stage of the tide, the time length of exposure
of the tidal flat within a tidal cycle or between spring tides
and neap tide phases, and the salinity of the source water.
The salinity of the tidal flat groundwater tends to be rela-
tively constant from tide to tide during a given season
and can even be constant over the entire year. However,
depending on the style of recharge into the pore waters
by flooding surface tides and a water table rising with
a tide, the salinity of the groundwater of estuarine tidal
flats can change seasonally. Although the water table of
the groundwater under tidal flats is shallow, it is separated
from the surface by a temporary vadose zone, and at low
tide generally, it is not in direct contact with the agencies
of solar-induced evaporation and wind shear. During
a flooding tide, the groundwater by rising into the tempo-
rary vadose zone salinized during the previous low-tide
period redissolves salt, hence raising its salinity. Plants
on the tidal flat, while clearly utilizing pellicular water in
 TIDAL FLAT SALINITY GRADIENT
Tidal Flat Salinity Gradient, Figure 1 (a) The characteristics of the water residing under tidal flats during high and low tide. The
components of the tidal flat hydrology during low tide are annotated in inset (b).
the temporary vadose zone, also utilize waters of the phre-
atic zone. Transpiration by tidal flat vegetation increases
the salinity of the pellicular water and the groundwater.
The temporary vadose zone is wetted and dried on
a tidal cycle and is subject to more direct evaporative pro-
cesses, causing the salinity of its pellicular water to be
more variable, to fluctuate over a tidal cycle, and to vary
from spring tide to neap tide and between the seasons.
Controls of tidal flat salinity
There are a number of factors that control local tidal
flat salinity and tidal flat salinity gradients. These are as
follows: (1) recharge of tidal flat pore waters and pellicular
waters by tidal inundation, (2) decreasing inundation of
the tidal flat upslope and increasing exposure to evapora-
tion by solar radiation and wind, (3) transpiration, (4) the
tidal range and nature of the tide, (5) the tidal flat slope,
(6) sediment types, (7) shoaling sediment bodies,
(8) homogeneity or heterogeneity of the tidal flat stratigra-
phy, and (9) freshwater seepage onto tidal flats from the
hinterland.
Specifically for tidal flats within estuaries, the salinity
of the open estuarine waters that daily recharge/inundate
the tidal flats and recharge the pore waters and pellicular
waters is a large factor in influencing tidal flat salinity as
this determines the starting salinity of the tidal flats that
will be subject to evaporative concentration.
Tidal flats facing open estuarine waters that are near the
marine realm and predominantly of marine salinity will be
recharged by waters of salinity
35 ppt, and evaporative
concentration will begin from that salinity. A fourfold
increase in salinity in higher parts of such tidal flats may
result in a salinity of
140 ppt. Tidal flats facing open
estuarine waters that are predominantly brackish (for
instance, in mid-estuarine locations) will be recharged by
waters with, say,
10–20 ppt salinity; a fourfold increase
679
in salinity in higher parts of the tidal flat in such situations
may result in a salinity of
40–80 ppt. Tidal flats facing
open estuarine waters that are predominantly freshwater
(for instance, in riverine and near-riverine locations
in the upper estuary) will be recharged by waters of
much lower salinity, and a fourfold increase in salinity
in higher parts of the tidal flat may result in a salinity still
in the freshwater field or lower brackish water field.
In each case, the tidal flat pore water and pellicular water
will begin their evaporative concentration from the
level of the salinity of the open waters that recharge the
tidal flats.
The topography of the tidal flat sloping from low tide to
high tide is a gradient of decreasing inundation. As such,
the progressively higher parts of a tidal flat are exposed
to increasing evaporation. The effects of solar radiation
and wind are direct, as they interface with large and exten-
sive wet or moist tidal flat surfaces. As water is evaporated
from the tidal flat surface (thereby concentrating salts or
even precipitating salts) and surface sediment begins to
dry, capillary action can deliver pellicular water to the sur-
face from the shallow water table underlying the tidal flat
either incorporating pellicular water with concentrated
salts or dissolving the precipitated salts. Both processes
increase salinity.
Transpiration by tidal flat vegetation such as man-
groves, saltmarsh, samphires, and seagrass also results in
moisture loss from tidal flats and in the concentration of
salts. Vegetation type on the tidal flat and its transpiration
rate determine the extent of moisture loss and the increase
in salinity.
The tidal range and the nature of tide (whether it is diur-
nal, semidiurnal, or mixed) determine the period that
a tidal flat is exposed and the area of exposure. Microtidal
coasts (say, with tidal ranges <0.5 m) tend to have
relatively narrow tidal flats and the exposure gradient
between low and high water results in relatively narrow
680 TIDAL FLAT SALINITY GRADIENT
shore-parallel zones of differing exposure indices that
increase towards the shore. Macrotidal coasts (say, with
tidal ranges >4 m, and locally over 10 m) tend to have rel-
atively wide tidal flats, particularly if the tidal flats have
a very low topographic gradient, and the exposure
between low and high water results in wide shore-parallel
zones within a tidal flat of differing exposure indices
increasing towards the shore. In this latter situation, the
gradient of salinity of the tidal flat is markedly evident
between MSL and high water and follows the shore-
parallel exposure indices.
In combination with sediment type, the tidal flat slope
determines how rapidly during a low tide the tidal flat
drains by vertical infiltration or by direct runoff. Tidal flat
slope also determine how rapidly tidal flat water changes
from phreatic (with its water table at, or close to the tidal
flat surface) to vadose (the wetted sediment above
a falling water table at or close to the tidal flat surface).
Topographically very low-gradient tidal flats, particularly
if they are muddy and not sandy, drain water relatively
slowly such that there is a longer residency time of pore
water and pellicular water in the surface and hence
a longer period for evaporation. Steeper flats, particularly
if they are sandy, tend to drain water more quickly with
shorter residency time for surface pore water and
pellicular water and less time for direct evaporation.
Sediment types also determine how rapidly the tidal flat
drains during low tide. Sandy substrates are more hydrau-
lically transmissive than mud substrates and drain water
more rapidly laterally and vertically. Because of their
low transmissivity, surface layers of mud and muddy sand
retain water for a longer period, are subject to evaporative
processes for longer periods, and tend to salinize more
rapidly than sand. While tidal flats that are composed
wholly of mud or wholly of sand will manifest relatively
uniform but different shore-parallel gradients of salinity
in groundwater and pellicular water in response to inunda-
tion, evaporation and transpiration, tidal flat sediments
that vary from sand to mud in cross-tidal-flat gradients,
or mosaics of sand and mud will exhibit more complicated
salinity patterns as the sediment types become additional
controlling factors.
The shore-parallel zones of increasing groundwater and
pellicular water salinity from MSL to high water in
response to decreasing inundation and increasing evapora-
tion and transpiration and their resultant salinity gradients
can be disrupted by the hydrological changes effected by
vertical sedimentary shoaling to levels above the high
water mark and by discordant (or heterogeneous) stratigra-
phy such as sandbars and cheniers. In humid environ-
ments with excess rainfall, the sedimentary sequence
may vertically shoal (accrete) to levels of high water and
then be capped by peat (Coleman et al., 1970). Peat,
sustained by rainfall, may continue to accrete vertically
to form peat mounds to levels above high tide. Such peat
mounds, being above the level of high tides, are
replenished by rainfall and store freshwater as subsurface
mounds which discharge into the adjoining tidal flats
diluting and perturbating the pattern of tidal flat salinity.
Similarly, a discordant body of topographically emergent
sand on a muddy tidal flat (e.g., a sandy chenier that has
emerged above high water) will not be hydrologically
and hydrochemically influenced by flood-tide recharge
but by rainfall. The chenier, on receiving rainfall, stores
freshwater as a subsurface lens which, during times of
elevated water tables after rain, can discharge into the sur-
rounding tidal flat, perturbating the salinity gradient (i.e.,
the increase in salinity from MSL to high water on the sur-
rounding tidal flat). The extent of the perturbation/dilution
by the freshwater stored in a chenier on the salinity of the
tidal flat depends on where the chenier is located. The
most profound effect is by cheniers and their freshwater
lenses on the higher parts of tidal flats where there is the
most marked contrast between freshwater and the tidal flat
salinity.
As such, tidal flats generally have a gradient of increas-
ing salinity from low-tidal zones to high-tidal zones
(Figure 2).
While there may be a general trend of increasing tidal
flat salinity from low-tidal levels to the high-tide mark,
this can be reversed by freshwater seepage along the mar-
gin of the hinterland. Freshwater seepage can manifest as
discharge along the upper tidal flat surface as surface
flows or springs or as subterranean flow along appropriate
stratigraphic conduits or sheets.
Tidal flat salinity gradients
Examples are provided of tidal flat salinity gradients in
Figures 3 and 4 from a tropical macrotidal estuarine
muddy tidal flat where rainfall is
1,548 mm per annum
and evaporation is
2,800 mm per annum and from sub-
tropical (near-temperate) microtidal estuarine sand-and-
mud tidal flat where rainfall is
880 mm per annum and
evaporation is
1,300 mm per annum (Figure 4). In both
cases the open estuarine waters are near marine in salinity.
In the tropical macrotidal estuarine example, the tidal flat
salinity increases from 35 to 40 ppt at MSL to
>180
near the level of the highest astronomical tide (HAT). In
the subtropical microtidal estuarine example, the tidal flat
salinity increases from 35 ppt at MSL to
60 ppt at mean
high water spring tide (MHWS), but is diluted by freshwa-
ter seepage along the hinterland edge (see below). In areas
of seasonal high rainfall, and where the tidal flats are
underlain by sand and bordered by a hinterland of sand,
the hinterland may be an aquifer storing and seasonally
discharging freshwater by seepage into the tidal flat –
while there is a salinity gradient from low-tidal to high-
tidal level, it seasonally changes from saline to brackish
in the higher parts of the tidal flat over the dry to wet
season.
In estuaries where there is a strong seasonal to perennial
delivery of freshwater by surface seepage or by subterra-
nean seepage from supratidal environments (Semeniuk,
1983; Cresswell, 2000), the gradient of increasing salinity
upslope can be locally reversed from hypersaline to
 TIDAL FLAT SALINITY GRADIENT 681

Tidal Flat Salinity Gradient, Figure 2 Salinity of groundwater and pellicular water across a tropical tidal flat in northwestern
Australia. The salinity of the groundwater and pellicular increases from the low-tidal zone to the high-tidal zone, with the
groundwater being of a slightly higher salinity than the pellicular water. There is no freshwater seepage in this location, and so
the hypersalinity of the high-tidal flats is not diluted (compare with high-tidal salinity in Figures 3 and 4).

Tidal Flat Salinity Gradient, Figure 3 Salinity of groundwater across two transects of a tropical macrotidal tidal flat from the Lawley
River delta of northwestern Australia (modified from Semeniuk, 1983). Along both transects, the salinity of the groundwater markedly
increases from the level of mean high water neap tide (MHWN) to the level of the highest astronomical tide (HAT). Transect 1 is in an
area with minimal freshwater seepage. Transect 2 is in an area with marked freshwater seepage, and as such, hypersalinity of
groundwater is diluted along the hinterland edge.
682 TIDAL FLAT SALINITY GRADIENT

Tidal Flat Salinity Gradient, Figure 4 Annotated profiles of the salinity of groundwater across two transects along a subtropical
(near-temperate) microtidal tidal flat from the Leschenault Inlet Estuary, southwestern Australia (Modified from information in
Cresswell (2000) and Semeniuk et al. (2000)). Along both transects, the salinity of the groundwater increases from the low-tidal level
to the level of mean high water spring tide (MHWS). One transect borders a high dune, while the other borders a low dune. Both sites
exhibit the effects of freshwater seepage from the dunes to the tidal flats.

brackish or to freshwater at the contact of the tidal flat with In the latter case, calcite grains may alter to dolomite
the supratidal zone. Examples of the effect of freshwater (Scoffin, 1987; Semeniuk, 2010).
seepage on tidal flat salinity are shown in Figures 3 and 4.

Consequences of tidal flat salinity gradients Discussion and summary

The main consequences of tidal flat salinity gradients are
twofold: species zonation and mineral precipitation. The
main effect of a tidal flat salinity gradient is on the occur-
rence and survivorship of various biota resulting in zona-
tion of species across a tidal flat. The best examples of
zonation are afforded by mangroves and salt marsh
(MacNae, 1968; Tomlinson, 1986; Pennings et al., 2005;
Silvestri et al., 2005) as they are conspicuous with differ-
ent species visually distinct in terms of physiognomy,
foliage color, and height. As such, their zonation is com-
monly evident as vegetation banding. The benthos of tidal
flats also exhibit zonation, but here, since many such
organisms are infaunal, the zonation needs to be brought
out by sampling and mapping.
The other effect of a tidal flat salinity gradient is
the progressive precipitation of minerals or the
geochemical alteration of minerals. As pore waters and
pellicular waters increase in salinity, precipitation of car-
bonate and sulfate minerals may occur (Logan, 1974).
Calcite or aragonite may be precipitated at salinities
>70 ppt, and gypsum may be precipitated at salinities
>120 ppt. These minerals are precipitated interstitially
(cementing the tidal flat sediments) or as nodules. Where
there is freshwater seepage into the upper tidal flat, car-
bonate grain precipitation, carbonate grain dissolution,
and (geochemical) carbonate grain alteration may occur.
The prevailing pattern of tidal flat salinity across a tidal
flat is one of increasing groundwater and pellicular water
salinity upslope in response to decreasing inundation and
increasing effects of evaporation and transpiration. For
tidal flats recharged by marine water, the general trend is
for the salinity to progressively increase from
35 to
40 ppt between low water and MSL to in excess of
150 ppt at levels of high water. The magnitude and extent
of increasing salinity upslope is determined by climate,
which influences the amount of evaporation and precipita-
tion, the salinity of the source water recharging the tidal
flats, and the sediment types. Perturbations to a simple
increasing upslope salinity gradient can result from
mosaics of sediment types, such as sand patches amongst
mud flats, with their potentially different water-holding
capacity and drainage rates, and can result from the hetero-
geneous discordant emergent sediment bodies (such as
cheniers and peat mounds) within tidal flat systems which
can store and discharge freshwater onto/into the tidal flat
resulting in local pockets of lower salinity. Freshwater
seepage from the supratidal zone onto/into the tidal flat
can have a marked influence on the salinity gradient and,
through its diluting effect, can reverse the salinity gradient
that was increasing from MSL to landward. Freshwater
seepage affecting tidal flat salinity gradients is most
marked in high-rainfall areas.
 TIDAL FRESHWATER HABITAT
Bibliography
Coleman, J. M., Gagliano, S. M., and Smith, W. G., 1970. Sedimen-
tation in a Malaysian high tide tropical delta. In Morgan, J. P.
(ed.), Deltaic Sedimentation – Modern and Ancient. Tulsa: Soci-
ety of Economic Paleontologists and Mineralogists Special Pub-
lication, Vol. 15, pp. 185–197.
Cresswell, I. D., 2000. Ecological significance of freshwater seeps
along the western shore of the Leschenault Inlet estuary. Journal
of the Royal Society of Western Australia: Special Issue on the
Leschenault Inlet Estuary, 83, 285–292.
Logan, B. W., 1974. Inventory of diagenesis in Holocene-Recent
carbonate sediments, Shark Bay, Western Australia. In Logan,
B. W. (ed.), Evolution and Diagenesis of Quaternary Carbonate
Sequences, Shark Bay, Western Australia. Tulsa: American
Association of Petroleum Geologists Memoir, Vol.
22, pp. 195–249.
MacNae, W., 1968. A general account pf the fauna and flora of man-
grove swamps and forests in the Indo-West-Pacific region.
Advances in Marine Biology, 6, 73–270.
Pennings, S. C., Grant, M.-B., and Bertness, M. D., 2005. Plant
zonation in low-latitude salt marshes: disentangling the roles of
flooding, salinity and competition. Journal of Ecology, 93,
159–167.
Scoffin, T. P., 1987. An Introduction to Carbonate Sediments and
Rocks. Glascow: Blackie & Sons, Bishopbriggs.
Semeniuk, V., 1983. Mangrove distribution in Northwestern Aus-
tralia in relationship to freshwater seepage. Vegetatio, 53, 11–31.
Semeniuk, V., 2010. A note on calcite precipitates as encrustations
around sea rush roots and as microlaminae in high tidal zones
of western Leschenault Inlet estuary. Journal of the Royal Soci-
ety of Western Australia, 93, 195–199.
Semeniuk, V., Tauss, C., and Unno, J., 2000. Avicennia marina in
the Leschenault Inlet area. Journal of the Royal Society of West-
ern Australia: Special Issue on the Leschenault Inlet Estuary, 83,
317–334.
Silvestri, S., Defina, A., and Marani, M., 2005. Tidal regime, salin-
ity and salt marsh plant zonation. Estuarine Coastal Shelf Sci-
ence, 62, 119–130, doi:10.1016/j.ecss.2004.08.010.
Tomlinson, P. B., 1986. The Botany of Mangroves. Cambridge:
Cambridge University Press.
Cross-references
Cheniers and Regressive Bedforms
Mangroves
Saltmarshes
TIDAL FRESHWATER HABITAT
Judith Z. Drexler
U.S. Geological Survey, California Water Science Center,
Sacramento, CA, USA
Synonyms
Tidal freshwater ecosystems; Tidal freshwater wetlands
Definition
Tidal freshwater habitat consists of wetland (See
Wetlands) ecosystems, which are located at the far
upstream end of estuaries and downstream from nontidal
freshwater ecosystems. The salinity of tidal freshwater
683
habitat is generally less than 0.5 parts per thousand (ppt).
A variety of animal species, including fish, amphibians,
reptiles, and particularly birds, live in tidal freshwater
habitat permanently or for at least part of the year.
Description
Tidal freshwater habitat is a transitional habitat type
found along the ecological continuum between tidal
saline wetlands and nontidal ecosystems in both tropical
and temperate coastal regions. Such habitat occupies the
upstream reaches of estuaries and large coastal wetland
complexes. For example, in the temperate zone, tidal
freshwater habitat can be found upstream from
salt marshes in estuaries along the West, Southeast, and
Gulf coasts of the United States (Odum et al., 1984;
Drexler et al., 2009b; Johnson et al., 2011). In the tropics,
an example of tidal freshwater habitat is palm forest,
which occurs upstream from mangrove ecosystems
(Ewel, 2010).
Tidal freshwater habitat may contain forest,
scrub-shrub, and herbaceous plant (marsh) communities.
Examples of forest types (swamps) are bald cypress
(Taxodium distichum) in the southeastern United States,
dragon’s blood (Pterocarpus officinalis) in the Caribbean
basin, and nipa palm (Nypa fruticans) along coastlines and
estuarine habitats of the Indian and Pacific Oceans
(Rivera-Ocasio et al., 2007; Ellison et al., 2010;
Dubenstein et al., 2013). Scrub-shrub wetlands typically
contain a variety of woody vegetation generally less than
5 m in height. Examples of scrub-shrub plant species
include smooth alder (Alnus serrulata), black willow
(Salix nigra), and marsh elder (Iva frutescens) in the
Southeast and Sitka willow (S. sitchensis), red osier dog-
wood (Cornus stolonifera), and hardhack steeplebush
(Spiraea douglasii) in the Pacific Northwest of the United
States (Thomas, 1984; Coulling, 2002). Herbaceous plant
communities, particularly those found in the higher
reaches of tidal freshwater marsh (Figure 1), tend to have
greater plant diversity than forest or scrub-shrub commu-
nities. Typical high marsh species in the United States
include annual plants, such as great ragweed (Ambrosia
trifida), smooth beggartick (Bidens laevis), and knotweed
(Polygonum spp.), and perennials, such as rice cutgrass
(Leersia oryzoides), green arrow arum (Peltandra
virginica), cattails (Typha spp.), giant cutgrass
(Zizaniopsis miliacea), and bulrushes (Schoenoplectus
spp.). In Europe, patch-forming perennials such as cattails
(Typha spp.), bulrushes (Schoenoplectus spp.), reed
canary grass (Phalaris arundinacea), valerian (Valeriana
officinalis), and bur-reed (Sparganium spp.) tend to dom-
inate the high marsh (Whigham et al., 2009; Barendregt
and Swarth, 2013).
The physical conditions in tidal freshwater habitat
reflect its transitional role in the landscape. The salinity
of channels is usually less than 0.5 ppt. Tidal range (See
Tidal Ranges) is typically from < 0.5 to 2 m and can
potentially be greater upstream than downstream due to
684 TIDAL FRESHWATER HABITAT
Tidal Freshwater Habitat, Figure 1 An example of a tidal freshwater marsh in coastal South Carolina, USA.
constricting morphology that results in amplification of
the tides (Mitsch and Gosselink, 2000). In some places,
such as the Gulf Coast of the United States, wind-derived
tides with irregular patterns of flooding and drying are
much more important than lunar tides (Mitsch and
Gosselink, 2000). Mature tidal freshwater habitat may
contain thick, highly organic peat (See Peat) soils, which
have accreted over hundreds or thousands of years
(Drexler et al., 2009a, b).
A variety of animals, including fish, amphibians,
reptiles, and waterfowl, live in tidal freshwater habitat per-
manently or for at least part of the year. With regard to fish,
many spawn and spend their entire lives in channels
within tidal freshwater habitat. Ictalurids (catfish), cypri-
nids (minnows, shiners, and carp), and centrarchids
(sunfish, crappies, and bass) are three important families
of such fishes. Many centrarchids, such as the largemouth
bass (Micropterus salmoides) and sunfishes (Lepomis
spp.), are important sport fish (Odum et al., 1984). Other
estuarine fishes such as killifishes (Fundulus spp.) and
juvenile naked gobies (Gobiosoma bosci) are part-time
residents, using tidal freshwater habitat for a source of
food or nursery grounds. A number of anadromous (See
Anadromous) and semianadromous fishes (which live as
adults in the ocean or lower estuary, respectively) travel
through tidal freshwater habitat on their way to streams
to spawn. The juveniles use tidal freshwater habitat as
nursery grounds. In the Pacific Northwest, several threat-
ened and endangered salmonid species, including the
Upper Columbia River Spring Chinook salmon
(Oncorhynchus tshawytscha), have been shown to rely
on tidal freshwater habitat for this purpose (Johnson
et al., 2011).
In addition to fish, a variety of amphibians and reptiles
depend on tidal freshwater habitat. River turtles and water
snakes are common inhabitants, but perhaps the most con-
spicuous residents are American alligators (Alligator
mississippiensis), which are distributed throughout the
southeastern United States (Mitsch and Gosselink,
2000). American crocodiles (Crocodylus acutus) may also
be found in tidal freshwater habitat but are more likely to
inhabit brackish and saline environments in their range,
which extends from southern Florida to South America
(Cherkiss et al., 2011). The much larger and broad-ranging
saltwater crocodile (Crocodylus porosus) is distributed
mainly throughout Southeast Asia and Australia and can
be found in tidal freshwater habitat, particularly during
the wet season (Campbell et al., 2013).
A number of mammals make their home in tidal fresh-
water habitat. Some of these inhabitants are particularly
well adapted to wet environments such as beaver (Castor
canadensis), marsh rabbit (Sylvilagus palustris), marsh
rice rat (Oryzomys palustris), river otter (Lontra
canadensis), muskrat (Ondatra zibethicus), mink
(Neovison vison), and nutria (Myocastor coypus; an inva-
sive species (See Invasive Species) originally from South
America) (Odum et al., 1984). Other mammals, including
white-tailed deer (Odocoileus virginianus), opossum
(Didelphis virginiana), and raccoon (Procyon lotor) may
use tidal freshwater habitat to fulfill at least part of their
food or habitat requirements (Odum et al., 1984).
The diversity of plants in tidal freshwater habitat pro-
vides a range of ecological niches (See Ecological Niche)
highly suitable for a large number of resident and migra-
tory bird species. In fact, because of the broad range of
plant assemblages, tidal freshwater habitat supports some
 TIDAL FRESHWATER HABITAT
of the largest and most diverse bird populations of all wet-
land types. In a survey of tidal freshwater marshes, Odum
et al. (1984) counted 280 species of birds in a broad range
of sub-habitats. Shorebirds, including sandpipers, killdeer,
woodcocks, dunlins, as well as rails, use low marsh and
exposed mudflats. Seed-eating species such as
red-winged blackbirds, sparrows, and bobolinks use the
mid- and high marsh, which contains grasses and sedges.
Herons, egrets, ibises, bitterns, and other wading birds
depend on ponded areas and channels. The marsh plain
and open water areas are prime habitat for ducks, geese,
swans, and other waterfowl, which have been hunted by
human populations for thousands of years (Barendregt
and Swarth, 2013). Shrubs and trees are used by arboreal
birds such as flycatchers and swallows. Tidal marsh
habitat also provides hunting grounds for hawks, falcons,
eagles, owls, and other birds of prey.
Today, tidal freshwater habitat is recognized for provid-
ing important ecosystem services such as biodiversity sup-
port and water purification (Barendregt and Swarth,
2013). Despite this, much tidal freshwater habitat has been
transformed or lost, due to its close proximity to human
populations. Extensive regions containing tidal freshwater
habitat have been drained and reclaimed for agriculture
(Penland and Ramsey, 1990; Ibanez et al., 1997; Drexler
et al., 2009a). What remains has been impacted by nutrient
loading, contaminants, and/or hydrologic diversions
(Barendregt and Swarth, 2013). Sea-level rise presents
a further threat to tidal freshwater habitat, particularly
from changes in hydrology and increases in salinity, which
may dramatically alter plant communities (Barendregt and
Swarth, 2013). In suitable environments, wetland
restoration may be used to reverse this trend and regain
habitat values and ecosystem services provided by tidal
freshwater habitat.
Summary
Tidal freshwater habitat consists of wetland ecosystems
containing a wide variety of plant communities, a high
diversity of birds, and plenty of other wildlife adapted to
wet conditions. Such ecosystems are typically found at
the far upstream end of estuaries. This position in the land-
scape makes tidal freshwater habitat vulnerable to local-
ized impacts such as drainage, water diversion, and
pollution as well as regional and global processes such
as sea-level rise. In suitable environments, restoration of
wetlands may be used to regain ecosystem services and
habitat values provided by tidal freshwater habitat.
Bibliography
Barendregt, A., and Swarth, C. W., 2013. Tidal freshwater wetlands:
variation and change. Estuaries and Coasts, 36, 445–456.
Campbell, H. A., Dwyer, R. G., Irwin, T. R., and Franklin, C. E.,
2013. Home range utilisation and long-range movement of estu-
arine crocodiles during the breeding and nesting Season. PLoS
ONE, 8(5), e62127, doi:10.1371/journal.pone.0062127.
685
Cherkiss, M. S., Romañach, S. S., and Mazzotti, F. J., 2011.
The American crocodile in Biscayne Bay, Florida. Estuaries
and Coasts, 34, 529–535.
Coulling, P. P., 2002. A preliminary classification of tidal marsh,
shrub swamp, and hardwood swamp vegetation and assorted
non-tidal, chiefly non-maritime, herbaceous wetland communi-
ties of the Virginia Coastal Plain. Virginia Department of
Conservation and Recreation, Division of Natural Heritage.
Natural Heritage Technical Report 02-18.
Drexler, J. Z., de Fontaine, C. S., and Deverel, S. J., 2009a. The
legacy of wetland drainage on the remaining peat in the
Sacramento–San Joaquin Delta, California, USA. Wetlands,
29, 372–386.
Drexler, J. Z., de Fontaine, C. S., and Brown, T. A., 2009b. Peat
accretion histories during the past 6,000 years in marshes of
the Sacramento–San Joaquin Delta, CA, USA. Estuaries and
Coasts, 32, 871–892.
Dubenstein, J. A., Conner, W. H., and Krauss, K. W., 2013. Woody
vegetation communities of tidal freshwater swamps in South
Carolina, Georgia, and Florida (US) with comparisons to similar
systems in the US and South America. Journal of Vegetation
Science, doi:10.1111/jvs.12115.
Ellison, J., Koedam, N. E., Wang, Y., Primavera, J., Jin Eong, O.,
Wan-Hong Yong, J., and Ngoc Nam, V., 2010. Nypa fruticans.
In IUCN 2012. IUCN Red List of Threatened Species. Version
2012.2. www.iucnredlist.org.
Ewel, K. C., 2010. Appreciating tropical coastal wetlands from
a landscape perspective. Frontiers in Ecology and the Environ-
ment, 8(1), 20–26.
Ibanez, C., Antoni, C., Day, J. W., Jr., and Curco, A., 1997. Morpho-
logic development, relative sea level rise and sustainable
management and sediment in the Ebre Delta, Spain. Journal of
Coastal Conservation, 3, 191–202.
Johnson, G. E., Storch, A. J., Skalski, J. R., Bryson, A. J.,
Mallette, C., Borde, A. B., Van Dyke, E. S., and Sobocinski,
K. L., 2011. Ecology of juvenile salmon in shallow tidal fresh-
water habitats of the Lower Columbia River, 2007–2010. Final
Report: PNNL-20083. Prepared for the Bonneville Power
Administration under U.S. Department of Energy Contract
DE-AC05-76RL01830.
Mitsch, W. J., and Gosselink, J. G., 2000. Wetlands. New York:
John Wiley & Sons.
Odum, W. E., Smith, T. J., Hoover, J. K., and McIvor, C. C., 1984.
The ecology of tidal freshwater marshes of the United States east
coast: a community profile. U.S. Fish Wildlife Service
WS/OBS-83/27.
Penland, S., and Ramsey, K. E., 1990. Relative sea-level rise in
Louisiana and the Gulf of Mexico: 1908–1988. Journal of
Coastal Research, 6, 323–342.
Rivera-Ocasio, E., Aide, T. M., and Rios-Lopez, N., 2007. The effects
of salinity on the dynamics of a Pterocarpus officinalis forest stand
in Puerto Rico. Journal of Tropical Ecology, 23, 559–568.
Thomas, D. W., 1984. The vascular flora of the Columbia River
estuary. Wassman Journal of Biology, 42, 92–106.
Whigham, D. F., Baldwin, A. H., and Barendregt, A., 2009. Tidal
freshwater wetlands. In Perillo, G. M. E., Wolanski, E., Cahoon,
D. R., and Brinson, M. M. (eds.), Coastal Wetlands: An Integrated
Ecosystem Approach. Amsterdam: Elsevier, pp. 515–533.
Cross-references
Anadromous
Ecological Niche
Invasive Species
Peat
Tidal Ranges
Wetlands
686 TIDAL HYDRODYNAMICS
TIDAL HYDRODYNAMICS
Bruce Parker
Center for Maritime Systems/Davidson Laboratory,
Stevens Institute of Technology, Hoboken, NJ, USA
Definition
Tidal hydrodynamics refers to the physics of vertical and
horizontal water movement caused by the tidal forces of
the moon and sun.
Introduction
In most estuaries, the dominant forces that cause the water
to move are produced by the gravitational effects of the
moon and sun on the oceans. These astronomical forces
cause the water to move vertically (the tide) and horizon-
tally (the tidal current), the entire phenomenon usually
referred to as the tides. Tidal motion is actually the motion
of extremely long waves generated in the deep ocean,
which propagate over the shallower continental shelf and
up into even shallower estuaries where their size is usually
amplified.
Although it is astronomical factors that makes the tides
so predictable, it is the hydrodynamics (i.e., physics of the
water movement) that determines the size and timing of
the tides. It is the hydrodynamics (determined by the
dimensions of the ocean, continental shelves, bays, and
rivers) that determines how large the tide range (height dif-
ference between high water and low water) will be and
when the high and low waters will occur. It is the hydrody-
namics that determines how fast the tidal currents will
flow and when slack waters will occur, and it is the hydro-
dynamics that determines how significant the diurnal tidal
signal will be compared with the usually dominant semidi-
urnal tidal signal.
Only the oceans are large enough for the tide-
generating forces to directly produce a tide of significant
size (see Tides, and also Parker, 2007). The tides in an
estuary are forced at the estuary entrance by the tide wave
from the deep ocean (having been modified to some
degree by propagating over the shallower continental
shelf). When we speak of the tidal hydrodynamics of estu-
aries, we are essentially speaking of shallow-water tides,
which are significantly different than open-ocean tides in
a number of ways, including having much larger tidal
ranges and faster tidal currents. In shallow water the tidal
wavelength is shorter, which is one factor leading to larger
tidal amplitudes when the tide wave reflects from the head
of an estuary. In estuaries, full and partial reflections of
these very long tidal waves, the nearness of basin lengths
to resonance for particular tidal frequencies, continuity
effects, frictional damping, and advective/inertial effects
all affect the tide and tidal currents. In shallow water the
hydrodynamics also transfers tidal energy, through vari-
ous nonlinear processes, to new frequencies. These new,
so-called shallow-water tidal constituents can be larger
than many of the astronomically generated tidal
constituents. Some shallow-water constituents (called
overtides) are higher harmonics of particular astronomical
constituents, which distort the simple sinusoidal shape of
tide and tidal current curves that are found in the open
ocean, often causing asymmetry in the tidal cycle. These
same nonlinear processes also lead to interactions between
the tide and other nontidal phenomena such as wind-
induced changes (e.g., storm surges) and river discharge.
Brief astronomical background
Tidal hydrodynamics deals with the forced oscillations
caused by the astronomical tide-producing forces. The
tides are the most predictable phenomenon in estuaries
because of this astronomical forcing, which varies due to
the well-known periodic motions resulting from the revo-
lution of the Earth around the sun, the revolution of the
moon-Earth system, and the rotation of the Earth on its
axis (see Tides, and also Parker, 2007). These periodic
motions determine that in any physical data set from an
estuary (e.g., water level, currents, salinity, and water tem-
perature), the tidal energy will be found at numerous very
precisely defined tidal frequencies, whose amplitudes and
phases can easily be calculated by harmonically analyzing
that data set. There are dozens of these so-called tidal con-
stituents, each constituent representing some aspect of
the relative astronomical motions of the Earth, moon,
and sun. These constituents tend to fall into bands. In the
semidiurnal band, constituents produce two high waters
(and two low waters) per day. In the diurnal band, constit-
uents produce one high water (and one low water) per day.
There are also other less important bands, including higher
harmonics caused by shallow-water nonlinear effects,
which will be discussed later in this entry. Tidal hydrody-
namics modifies the amplitude and phases of these tidal
constituents.
For the purposes of this entry, only the five most impor-
tant tidal constituents will be mentioned until the shallow-
water constituents are discussed (for more constituents see
Parker, 2007, Table A, which includes 149 tidal constitu-
ents). These five larger constituents are named M2, S2,
N2, K1, and O1, the first three being semidiurnal and the
last two diurnal. M2, the main lunar semidiurnal constitu-
ent, represents the Earth turning under a slowly revolving
moon with 2 cycles (and two high waters) in each
24.8412-h lunar day. It thus has a period of 12.4206 h
and a frequency (1/tidal period) of 1.9323 cycles per
(solar) day. S2, the main solar semidiurnal constituent, rep-
resents the Earth turning under the sun with 2 cycles (and
two high waters) in each 24.00-h solar day. It thus has
a period of 12.00 h and a frequency of 2.00 cycles per
(solar) day. When the moon and sun are in alignment at
new and full moons, their tidal forces work together to
produce larger tide ranges (called spring tides). When
the moon and sun are out of alignment, at first and third
quarters, their tidal forces work against each other to pro-
duce smaller ranges (called neap tides). N2, with a period
of 12.6583 h, represents the effect of the elliptical
 TIDAL HYDRODYNAMICS
Earth–moon orbit, in which the distance between the
moon and Earth varies over a 27.5546-day period, from
perigee (the moon closest to the Earth, and so a stronger
tidal force) to apogee (the moon farthest from the Earth,
and so a weaker tidal force) and back to perigee. This mod-
ulates the main lunar tidal force (M2), and that modulation
is represented by N2 combined M2. The diurnal lunar tidal
forces resulting from lunar declination are represented by
two diurnal tidal constituents, O1 and K1, with periods of
25.8193 and 23.9345 h (and frequencies of 0.9295 cpd
and 1.0027 cpd). The minimum combined effect of these
two constituents occurs every 13.66 days, at the times
when the moon is over the equator. Their maximum com-
bined effect occurs at maximum lunar declination, when
the moon is either farthest north of the equator or farthest
south of the equator. Because of these well-known astro-
nomical frequencies, some form of tide prediction has
existed since even the earliest civilizations. For a history
of tide prediction, see Parker (2012).
Hydrodynamic effects on tide ranges and tidal
current speeds in estuaries
When the very long tide wave generated in the deep ocean
reaches the shallower water of the continental shelf and
the even shallower water of estuaries, it is slowed up,
amplified, modulated, and distorted by a number of hydro-
dynamic mechanisms. To understand what happens to
a tide wave in an estuary, it is helpful to first look at two
opposite extremes, the tide wave in a long river and the
tide wave in a closed-off bay, both basins being deep
enough for a minimal damping effect from bottom
friction.
Progressive tide waves
The tide wave propagates up the river as a progressive
wave (see Figure 1, top panel), which means that the crest
of the wave (high water) moves progressively up the
river, as does the trough of the wave (low water). In such
a progressive tide wave, the maximum flood current
(namely, when the current is flowing the fastest up the
river) occurs at the same time as high water, and the max-
imum ebb current (namely, when the current is flowing
the fastest down the river) occurs at the same time as
low water. Slack water (when the current speed is zero)
occurs exactly halfway between high water and low
water. If the river is of constant width, the amplitude
(tide range) of this progressive tide wave will not change
as it moves up the river. However, if the width decreases
as going upriver, then the amplitude (tide range) will
increase, because the same amount of water is being
forced through a smaller basin cross section. If the depth
of the river decreases, there is a similar though less dra-
matic amplifying effect (which is generally outweighed
by the increased energy loss from bottom friction due to
the shallower depths).
687
Standing tide waves
In a closed-off bay, the tide wave progressing up the bay
will be reflected at the closed end and will travel back
down the bay. This reflected wave is not observable by
someone on the shore because it is superimposed on the
incoming tide wave propagating up the bay, and it is
the combination of the two waves that is observed. The
resulting combined wave is called a standing wave,
because the high and low waters do not progress up the
bay (see Figure 1, bottom panel). The water surface simply
moves up and down everywhere at the same time, with the
largest tide range at the head of the bay. With a standing
tide wave, the tide range increases as one moves from
the ocean entrance toward the closed end of the bay,
assuming the length of the bay is less than or equal to
one fourth of a tidal wavelength from the head of the
bay. If the bay is longer than that, there will be a location
with minimum tide range (at one fourth of a tidal wave-
length from the head of the bay), so from the entrance
moving up the bay the tide range would first decrease
and then increase. This location is explained by looking
at the incident and reflected progressive waves. In a
progressive wave, high water comes one half a
wavelength before low water, so if the high water of
a progressive wave travels a distance equal to one fourth
of a tidal wavelength up the bay to the head, where it is
reflected, and then travels one fourth of a wavelength back
down the bay, it will have gone one half a wavelength and
so coincide with low water of the incoming progressive
wave, and the two will roughly cancel each other out at
that location, producing the minimum tide range (if there
was no friction and the incident and reflected wave were
equal in amplitude, the minimum range would be zero
and it would be called a node).
For a standing wave, high waters occur at the same time
everywhere on one side of the minimum (node), which is
the same time that low waters occur on the other side. The
strongest tidal currents occur when water level is near
mean tide level, halfway between the times of low water
and high water. At the times of high water and low water,
there is no current flow (slack water). The water flows into
the bay, stopping the inward flow at high water; reverses
direction; and flows out of the bay until low water, at
which time it reverses again and starts flowing into the
bay again.
Amplification of tides in an estuary
The tidal wavelength is determined by the depth of the
estuary. If friction is ignored, the formula for the tidal
wavelength, l, is l ¼ T(gD)½, where D is the depth of
the water, T is the tidal period, and g is the acceleration
due to gravity. The shallower the bay, the shorter is the
wavelength. When the length of the bay equals one fourth
of a tidal wavelength, then the bay’s natural period of
oscillation will be the same as the tidal period. One finds
the largest tide ranges in bays that are exactly one fourth
of a tidal wavelength long, due to what is called resonance.
688 TIDAL HYDRODYNAMICS

Tidal Hydrodynamics, Figure 1 (Top panel) An idealized frictionless tide wave propagating up a river as a progressive wave. High
water occurs later as one moves upstream. (Bottom panel) An idealized frictionless tide in a bay as a standing wave (the water level is
shown for two opposite extremes, high water and low water). High water occurs at the same time everywhere on one side of the
node (the point of zero range).

When the water in the bay is forced to move up and down
by the tide at the entrance, it will freely oscillate (slosh up
and down and back and forth) with a natural period that
depends directly on the bay’s length and inversely on the
square root of its depth, namely, T ¼ l/(gD)½. If the basin
has the right combination of length and depth so that the
natural period is exactly the same as the tidal period, then
the oscillation inside the bay will be synchronized with the
oscillation at the entrance due to the ocean tide. In other
words, the next ocean tide will be raising the water level
in the bay at the same time that it would already be rising
due to its natural oscillation (stimulated by the previous
ocean tide wave), so that both are working together, thus
making the tide range inside higher. (In the real world,
friction keeps these resonating oscillations from being
infinitely large, by taking away some of the energy.)
In the above discussion, bottom friction was considered
minimal because of deep water, but bottom friction greatly
affects all hydrodynamics and is especially important in
shallow waters. Because of bottom friction, the tide wave
in real estuaries actually falls in between the two extremes
of the pure progressive wave and pure standing wave
described above. This is because friction reduces the
amplitude of the tide wave as it travels. Thus, the reflected
wave will always be smaller than the incoming wave,
especially near the bay entrance (since the reflected wave
has traveled longer to get all the way back to the entrance),
and the combination of these two frictionally damped pro-
gressive waves will not be a pure standing wave. There
will be no point of zero tidal range (no node), but only
an area of minimum tidal range (a quasinode). There will
be some progression of high waters and low waters up
the bay, but not as quickly as a pure progressive wave.
This progression will be faster near the entrance and
slowest near the head of the bay. Maximum flood or ebb
currents will not occur exactly halfway between high
water and low water. A basin one fourth of a wavelength
long will still produce the largest possible tidal range at
 TIDAL HYDRODYNAMICS 689

4.0
heavily μ=8.0 μ=6.0 μ=5.0 μ=4.0
damped
progressive

RATIO OF ELEVATIONS AT HIGH WATER, ηH /η 0

wave

3.0

μ=3.0

2.0
μ=2.5

μ=2.0
μ=1.5
μ=1.0
1.0 μ=0.5

almost
standing
wave
0.0
−0⬚ −10⬚ −20⬚ −30⬚ −40⬚ −50⬚ −60⬚ −70⬚ −80⬚ −90⬚ −100⬚−110⬚−120⬚−130⬚−140⬚−150⬚−160⬚−170⬚−180⬚
CLOSED M PHASE DIFFERENCE RELATIVE TO REFLECTION POINT, kx
END OF 2
WATERWAY

Tidal Hydrodynamics, Figure 2 The amplitude ratio (¼tide range ratio) plotted versus distance from the closed end of the waterway
(extreme left of plot) in terms of wavenumber phase, kx, for several values of the frictional damping coefficient, m. kx ¼ 90o
represents one fourth of a tide wavelength from the closed end of the bay.

the head of the bay, but friction will keep that tide range little frictional damping (m ¼ 0.5), one sees an almost
much smaller than it would be without friction. standing wave with a nearly zero tide range at one fourth
(Mathematical formulas representing the effects described of a tide wavelength (kx ¼ 90o), that location being
above are derived in Parker, 2007). the quasinode. For the case of very large frictional
Figure 2 illustrates how friction affects the tide range damping (m ¼ 8.0), one sees an almost pure damped pro-
along a waterway. It is based on a relatively simple analyt- gressive wave that decreases in tide range as it moves up
ical model of an exponentially damped tide wave that the waterway. (For this last case, it could also represent
reflects off the closed end of the waterway (Parker, an open-ended river that is not very shallow, but for which
2007). In this figure, the ratio of the tide range (at any loca- there is no reflected wave because it is not closed off.)
tion) to the tide range at the closed end is plotted for vari- In general, one sees very large amplification for small m
ous values of the damping coefficient m. This is for waterway lengths that are equal to or less than one
a convenient way to put many cases on the same plot, fourth of a tide wavelength (kx ¼ 90 ). For
because every case can be plotted relative to the closed m ¼ 0 (no friction), there is infinite amplification at exactly
end. (Such things as amplification are easier to visualize one fourth of a wavelength (namely, resonance). That
if values are plotted relative to the entrance, but then each never happens in the real world (because of friction), but
case would have to be plotted separately.) In Figure 2, if very large tide ranges do occur in bays with lengths that
one imagines the entrance at, for example, one fourth of are near a quarter tidal wavelength. For waterways that
a tide wavelength from the closed end of the estuary are shorter or longer than one fourth of a wavelength, the
(indicated by kx ¼ 90o in the figure), then one can see amplification is not as great (for the small m case; for the
that for m ¼ 0.5 (which represents very little frictional large m case there will never be amplification because of
damping in a deep waterway) the tide range increases sig- the strong frictional damping). From this figure, one can
nificantly as one moves up the estuary toward the closed see that for a deep waterway (small m) that is one half
end (in the figure, moving to the left from kx ¼ 90o). a tidal wavelength long (kx ¼ 180o), the tide range
At the other extreme, for m ¼ 8.0 (great frictional damping decreases until one reaches one fourth a wavelength and
in a very shallow estuary) the tide range decreases signif- then increases until at the closed end of the waterway the
icantly as one moves up the estuary. For the case of very tide range is almost the same as it was at the entrance.
690 TIDAL HYDRODYNAMICS
The largest tides and fastest tidal currents
The largest tide ranges are found in bays that are close
to one fourth of a tidal wavelength long. Tide ranges
reach 15 m (50 ft) in Minas Basin in the Bay of Fundy
and in Ungava Bay (also in Canada). Tidal ranges
greater than 12 m (40 ft) occur at the northern end of
Cook Inlet near Anchorage in Alaska, in Bristol Bay
in the United Kingdom, in the Magellan Strait in Chile,
in the Gulf of Cambay in India, and along the Gulf of
St. Malo portion of the French coast bordering the
English Channel. In some bays, the very high tide range
at the head of the bay is due to a combination of both
a narrowing width and a near resonant situation (due
to the right length and depth). The highest tide ranges
may involve several amplifications, the bay being per-
haps connected to a gulf which is perhaps connected
to a wide continental shelf, with amplifications of the
tide wave occurring in each basin. This is the case with
the Bay of Fundy tides, the tide wave being already
amplified by the continental shelf and the Gulf of Maine
prior to entering the Bay of Fundy. Huge tide ranges
are not restricted to bays. If the continental shelf is the
right combination of depth and width, a near resonant
situation can also result. This is the reason for the
12-m (40-ft) tidal ranges along the coast of southern
Argentina. The continental shelf there is over 965 km
(600 mi) wide and includes the Falkland Islands near
the edge of the shelf, where the tide range only reaches
2.0 m (6.5 ft). The distance from the Argentinean coast
to the edge of the shelf is fairly close to one fourth of
a tidal wavelength for that depth of water. Essentially,
that wide shelf has a natural period of oscillation that
is fairly close to the tidal period.
The largest tidal currents in estuaries tend to be near the
entrances. (Maximum tidal current speeds are zero at the
head of the bay, since there is no place for the water to
flow). As one moves down an estuary toward the ocean,
the maximum flood and maximum ebb tidal current
speeds increase, with the greatest speeds occurring at the
entrance, or, if the estuary is long enough, at the area of
smallest tide range (the nodal area). However, if the width
of the estuary decreases at any point, the current speeds
will be increased in that narrow region. (It must flow
faster, since the same volume of water is being forced to
flow through a smaller cross section). This can be espe-
cially dramatic if there is a sudden decrease in width and
depth. The largest tidal currents are found in narrow straits
in which the tides at either end have different ranges or
times of high water. Tidal current speeds greater than
7.7 m/s (15 knots) occur in Seymour Narrows, between
Vancouver Island and the mainland of British Columbia,
Canada. Tidal currents of 5.1 m/s (10 knots) are found in
South Indian Pass in Southeast Alaska and in Kanmon
Strait, Japan. Fast tidal currents in narrow straits that sud-
denly widen can also produce a tidal whirlpool, a violently
rotating funnel-shaped hole of water, which is explained
later in this entry.
Coriolis effects on tide ranges in wide estuaries
If an estuary is wide enough, one also sees larger tide
ranges on the right side of the bay (looking up the bay)
due to the Coriolis force. The Coriolis force is
a fictitious force due to our observing motion from the
rotating reference frame of the Earth (see Parker, 1998).
It acts perpendicular to the flow of the water, thus pushing
water currents to the right in the Northern Hemisphere
(and to the left in the Southern Hemisphere) as they flow.
For a pure progressive tide wave (no friction), at high
water the tidal current flows up the bay, so the tidal height
will be greater on the right-hand shore than on the left-
hand shore. At low water, the tidal current flows down
the bay, so the tidal height will be lower on the right-hand
shore than on the left-hand shore. The result is that the tide
range (high water minus low water) will be greatest on the
right-hand shore (looking up the bay). For a pure standing
wave (no friction), the pattern of high water caused by the
Coriolis force is more complicated, as is shown in the
upper half of Figure 3. This figure shows lines of constant
tide range (corange lines), as well as lines of locations with
the same time of high water (cotidal lines), in an idealized
rectangular basin for the case where the effect of bottom
friction is ignored. A single point of zero tidal range
(a node) occurs in the center of the bay one fourth of
a tidal wavelength from the head of the bay.
This figure comes close to representing the corange and
cotidal lines in a bay that is very deep. A more typical case,
including the damping effect of bottom friction, is
shown in the bottom half of Figure 3. In this case the node
has moved to the left (when looking up the bay)
and becomes a virtual node since it is on land. (See
Parker, 2007, for derived mathematical formulas
describing this effect.) One can see some similarity
between the pattern of corange lines in the lower half of
Figure 3 and the M2 corange lines in Figure 4 for the
Strait of Juan de Fuca–Strait of Georgia (although the
latter has many geographic variations not included in the
simple regular basin of Figure 3). In Figure 4 the pattern
of a quasinode is also seen to the southwest of Victoria.
Hydrodynamic effects on diurnal versus
semidiurnal tides in estuaries
In many estuaries it is common to have semidiurnal tides,
namely, two high waters (and two low waters) a day, with
the heights of the two high waters (and two low waters)
being approximately the same size. In some estuaries
and seas, there can be diurnal tides, namely, only one high
water (and one low water) per day. Most common, how-
ever, is to have mixed tides, namely, two high waters
(and two low waters) a day but where there is considerable
difference between the heights of two consecutive high
waters (and/or between the heights of the two consecutive
low waters), that difference being due to a strong diurnal
signal. It is tidal hydrodynamics that determines which
of these three types of tide one will see at particular
 TIDAL HYDRODYNAMICS
Tidal Hydrodynamics, Figure 3 The effect of Coriolis force on the M2 tide range (corange lines) and the time of high water (cotidal
lines) for an idealized rectangular bay. The top panel shows the case with no bottom friction; the bottom panel includes the effect of
bottom friction.
locations in an estuary. This classification of tides is not
always as descriptively precise as one might wish, because
the relative strength of the diurnal signal to semidiurnal
signal varies throughout the month, with the diurnal signal
being strongest during maximum lunar declination (i.e.,
when the moon is farthest north or south of the equator).
Thus, a so-called mixed tide might have two high waters
a day when the moon is over the equator, but only one high
water a day when the moon is farthest north or south of the
equator (and thus being diurnal at those times).
The dimensions of a basin (and the dimensions of the
waterways leading to the basin) determine the size of the
diurnal tidal signal (primarily K1 and O1) compared with
the usually dominant semidiurnal tidal signal (primarily
M2, S2, and N2). A particular bay could have a natural
period of oscillation that is closer to the diurnal tidal
period (approximately 24.84 h) than to the semidiurnal
period, thus amplifying the diurnal forcing at the entrance
to the bay more than the semidiurnal signal. Depending on
the size of the diurnal signal at the entrance, the result
could be a mixed tide or a diurnal tide. At such locations
(e.g., parts of the Gulf of Mexico), the tide will be diurnal
near times of maximum lunar declination, but will be
mixed near times when the moon is over the equator.
As mentioned earlier, the wavelength, l, of a tide wave
in a bay depends on the depth of the water, D, and on
the tidal period, T, according to l ¼ T(gD)½ (if frictional
effects are ignored). The longer the tidal period, the longer
is the tidal wavelength. A diurnal tidal component has
691
a wavelength twice as long as a semidiurnal tidal compo-
nent, since its period is twice as long. When a waterway is
shallow enough and long enough so that more than one
fourth of a semidiurnal (e.g., M2) wavelength fits in the
waterway, there will be a nodal area with a very small
semidiurnal (M2) tidal range. This will be an area where
the diurnal tide could dominate, since the diurnal tide
would still be large at the semidiurnal nodal area (the diur-
nal node being twice as far from the head of the bay).
Thus, near the head of the waterway the tide could be
semidiurnal, but near the semidiurnal nodal area, the tide
could be mixed or even diurnal. This is the case near
Victoria, British Columbia, at the southeastern end of Van-
couver Island (see Figure 5). At that location along the
Strait of Georgia–Strait of Juan de Fuca waterway, the
M2 and S2 tidal constituents decrease to a minimum, but
the K1 and O1 constituents do not, and so the tide becomes
mixed mainly diurnal, while at the northern end of the
Strait of Georgia, the tide is mixed, mainly semidiurnal.
Whether due to a basin size conducive to amplifying
the diurnal signal or due to the existence of
a semidiurnal nodal area (leaving the diurnal signal as
the dominant one), there are numerous areas around the
world with strong diurnal tides – places like Norton Sound
in Alaska near the Bering Strait and various (but not all)
locations in the Philippines, New Guinea, and the islands
of Indonesia. In southern China, at Beihai, and at Do
Son, Vietnam, the diurnal signal is very dominant, with
diurnal tidal ranges that reach 4.6 m (15 ft) and 3.0 m
692 TIDAL HYDRODYNAMICS

Tidal Hydrodynamics, Figure 4 A corange chart showing the geographic variation of the M2 tide range for the Strait of Juan de
Fuca–Strait of Georgia. A minimum tidal M2 range occurs just to the southwest of Victoria, British Columbia (1 ft ¼ 0.305 m).

(10 ft), respectively (near times of maximum southern
declination of the moon). In these locations, the tide
remains diurnal even when the moon is over the equator.
Nonlinear effects of shallow water
on tides: overtides and compound tides
Shallow water also has other effects on the tide. It can, for
example, distort the shape of the tide wave, that is, make it
very asymmetric, so that its rise and fall (and its flood and
ebb) are no longer equal (see the second curve in Figure 6).
The tide can then no longer be described by a simple sine
wave (such as the first curve in Figure 6). In some cases,
such distortion leads to double high waters or double
low waters (see the third curve in Figure 6). The extreme
case of distortion is a tidal bore (the fourth curve in
Figure 6), when the tide wave becomes so steep that it is
essentially a continuously breaking wave moving up
a river as a turbulent wall of water.
Shallow water distorts the tide through several mecha-
nisms that are nonlinear – that is, in the equations of
motion (based on conservation of mass and momentum,
see Parker, 2007), each mechanism can be tied to
a specific term in which key parameters (such as water ele-
vation or velocity) multiply each other, which leads to
energy transfer. (Linear terms contain only one key param-
eter and thus their separate effects simply add, with no
interaction.) Such nonlinearities can only be handled ade-
quately in numerical hydrodynamic models (based on the
nonlinear equations of motion), rather than by simple
formulas from analytical models, such as those that we
used to described many of the linear tidal effects.
 TIDAL HYDRODYNAMICS 693

Tidal Hydrodynamics, Figure 5 Chart illustrating the geographic variation in the (K1 + O1)/(M2 + S2) tidal constituent amplitude ratio,
as well as the type of tide classification for each region in the Strait of Juan de Fuca–Strait of Georgia.

However, the effects of these nonlinear terms can be
explained physically here without resorting to the math
of these nonlinear models. (For mathematical treatments
of the nonlinear terms, see Parker, 2007).
The speed, C, at which a long tide wave travels depends
on the depth of the water, D, approximately (ignoring fric-
tion) as the formula C ¼ (gD)½. When the depth of the
water is much greater than the tidal range, the speed of
the crest of a tide wave and the speed of the trough are vir-
tually the same, since the tide wave itself has only a very
small effect on the total water depth. However, in the shal-
low water where the depth is not much greater than the tide
range, the total water depth under the crest is significantly
larger than the total water depth under the trough. In this
case, the crest of the wave (high water) travels faster than
the trough of the wave (low water). If the tide wave travels
far enough, the crest begins to catch up with the trough
ahead of it (which is falling behind the crest ahead of it).
Thus, high water arrives sooner than it would in deeper
water, and there is a faster rise to high water and
a slower fall to low water. The shape of the tide curve
could perhaps look like that shown in the second curve
in Figure 6. In terms of harmonic constituents, this distor-
tion transfers energy from M2 into the second harmonic,
a constituent called M4, with half the period of M2. Com-
bining an M2 tide curve and an M4 tide curve, one can pro-
duce the distorted tide curves shown in Figure 6, with the
M4/M2 ratio increasing as one goes from top curve to bot-
tom curve in those figures. The third curve in Figure 6
shows a double low water, but with a different phase rela-
tionship between M2 and M4, one could obtain a double
high water.
694 TIDAL HYDRODYNAMICS
Tidal Hydrodynamics, Figure 6 Typical tide curves (over one
and a half tidal cycles) for an area with (top panel) no shallow-
water effect and for three areas with increasing degrees of
distortion caused by the shallow water. The second panel shows
moderate distortion of the sinusoidal curve seen in the first
panel. The third panel shows a double low water. The fourth
panel shows the almost instantaneous rise in water level due to
the passage of a tidal bore.
Another shallow-water distorting mechanism is caused
by bottom friction, which can have both asymmetric and
symmetric effects. The asymmetric effect (similar to that
just discussed and represented in Figure 6) is produced
because friction has a greater effect in shallow water than
in deep water (there being less water to have to slow
down), and so it slows down the trough more than the
crest, contributing to the distortion of the tide wave and
the generation of M4. The symmetric effect is produced
because frictional energy loss is proportional to the square
of the current speed. This means that there will be much
more energy loss during times of maximum flood and
maximum ebb than near times of slack water
(or minimum flow). This results in the generation of
another higher harmonic, M6, with a period of one third
that of M2. This effect, combined with the asymmetric
effect, can lead to double high or low waters (such as the
third curve in Figure 6).
Higher harmonic tidal constituents like M4 and M6 are
referred to as overtides (a term analogous to the term over-
tones in acoustics). M4 is the first overtide (and the second
harmonic) of M2. M6 is the second overtide (and the third
harmonic) of M2. M8 is the third overtide (and the fourth
harmonic) of M2. Whereas M4 and M6 are generated by
first-order nonlinear processes, M8 is generated by second-
order nonlinear processes (i.e., it is generated by nonlinear
effects on M6).
Friction dissipates energy from the entire tide wave and
slowly wears the entire tide wave down. However, if, as
the tide wave propagates up the river, the river’s width is
decreasing significantly, this can keep the amplitude of the
wave high in spite of the friction. Thus, the tide wave can
continue to travel up a narrowing river, getting more and
more distorted in shape. A further distortion can be caused
by the river flow interacting with the tide (see below). In
the extreme case, the distortion from all these effects can
lead to the creation of a tidal bore, when the tide wave
becomes so steep that it is essentially a continuously break-
ing wave, moving up a river as a turbulent wall of water (see
fourth curve in Figure 6). (The earliest known tide table was
printed in 1056 AD for the tidal bore on the Qiantang River
in China; see Parker, 2012.)
New tidal constituents called compound tidal constitu-
ents can be produced through the nonlinear interaction of
two astronomical tidal constituents. For example, the
above symmetric quadratic friction effect causes the inter-
action of two tidal constituents, such as M2 and N2. M2
and N2 go in and out of phase over a 27.6-day cycle
(perigee to apogee to perigee). In this case the greatest
energy loss occurs when M2 and N2 are in phase and pro-
ducing the strongest tidal currents, and the lowest energy
loss occurs 13.8 days later when M2 and N2 are out of
phase and producing the weakest tidal currents. Because
energy loss is proportional to the square of the current
speed, the increased energy loss when M2 and N2 are in
phase is greater than the decreased energy loss when they
are out of phase, and the result is that each constituent will
be smaller than if it existed without the other present. The
 TIDAL HYDRODYNAMICS
Tidal Hydrodynamics, Figure 7 Water level data from the tide gauge at Trenton, NJ, during a high river discharge period (January
20–29, 1979). The tide range is reduced when the river discharge is high, and the tide curve is distorted (with a faster rise to high
water) (1 ft ¼ 0.305 m).
reduction in N2 (due to M2) will be greater than the reduc-
tion in M2 (due to N2), because M2 is much greater than
N2. However, M2 will be reduced by the combined
interactions of all the other tidal constituents. There is
a 27.6-day modulation of this energy loss from M2 and
N2, and this produces two new compound tidal constituents
called 2MN2 and 2NM2. (Similarly, the above asymmetric
mechanisms also cause interactions between constituents,
producing higher frequency constituents such as MN4 from
M2 and N2.) More compound tides and overtides and their
origin are explained by Parker (2007).
In shallow waterways with large tidal ranges, the
nonlinear effects can produce dozens of compound tides
and overtides of significant enough size that they must
be included in harmonically based tide predictions. For
example, for tide predictions at Anchorage, Alaska, at
the northern end of Cook Inlet, 114 tidal constituents
(most of them shallow-water constituents) must be used
to predict the 30-ft tides there (as compared with typically
26 or less constituents in other waterways).
Nonlinear tidal interaction with river flow
In a tidal river, water flow is due to both the tidal current
and the river current itself (i.e., the freshwater flowing
downhill). The result of the combined tidal current and
river current is a faster and longer-lasting ebb current
phase and a slower and shorter flood current phase. Far
enough up a river, where the river flow is faster than the
strongest tidal current, the flow of water will always be
downstream. In this case, the speed of flow will oscillate,
flowing the fastest downstream at the time when maxi-
mum ebb occurs farther down the river and flowing the
695
slowest downstream at the time when maximum flood
occurs farther down the river. This is a simple linear addi-
tion of the river current to the tidal current.
However, because of the shallow water, the river flow
also interacts with the tide nonlinearly and distorts it,
mainly due to the effect of bottom friction. Energy loss
due to friction is proportional to the square of the total
current speed. During ebb, the tidal current is in the same
direction as the river current, and the result is a larger
combined ebb current, with increased energy loss. Dur-
ing flood, the tidal current is in the opposite direction
as the river current and the result is a smaller combined
current, with reduced energy loss. This has an asymmet-
ric effect that distorts the tide (causing a faster rise to
high water, delaying the time of low water, and increas-
ing the size of M4). It also wears down the entire wave
further because the increased energy loss during ebb is
larger than the decreased energy loss during flood. In
Figure 7, one can see the tide range shrink when the river
discharge increases. The curves also become more asym-
metric, rising more quickly to high water and falling
more slowly to low water. A harmonic analysis of water
level data during this period of high river runoff would
give a smaller M2 and larger M4 than during times of
small runoff. (See Parker, 2007, for a mathematical treat-
ment of this effect and more examples.)
Another type of shallow-water effect causes interac-
tions between the tide and low-frequency storm surges
(generated by the wind) that have periods longer than tidal
periods. In this case, when the water level is raised by an
onshore wind, the water depth increases and changes the
tidal dynamics, usually increasing the tide range. When
an offshore wind lowers the water level, decreasing the
696 TIDAL HYDRODYNAMICS

North L+3 h
a LL+2
L+2 H-2
b LL+1h h
HH-3
h LL+3h
h HH-2
H-1 LL HL+2 h
L+1 h LH-2
h
HH-1
HL+1
LL-1h HH LH-1
h
H
L HL
h HH+1
h LH
LL-2

H+1 North LL-3h h

L-1 HL-1h HH+2 LH+1h
HH+3h
L-2 H+2 h
HL-2 LH+2h
H+3 1 knot LH+3h
1 knot

North d North
c
1.0 Knot 0.50 Knot
S+3.0
S+3.5
F+0.0 0.25
S+1.0
F+9.5

West East West East

1.0 0.25 S+0.0 S+4.0
Knot Knot
F+4.0
S+12.0

F+6.5 S+5.5
1.0 Knot
S+7.5
South

Tidal Hydrodynamics, Figure 8 Examples of three rotary tidal currents and one reversing tidal current (third example). The first two are
off an ocean coast and are idealized. The last two are within an estuary and are from actual data. For the first two examples, each vector
shows the speed and direction of flow for that hour of the tidal cycle, but for the last two examples only the tips of the vectors are shown.

water depth, the result is usually a decreased tidal range.
(See Tidal and Nontidal Oscillation; see also Parker,
2007, for a mathematical treatment of this interaction.)
Tidal effects on mean sea level: another
nonlinear effect
Tidal heights are referenced to some type of datum, typi-
cally the mean level of a particular key point on the tide
curve. Datums such as mean lower low water (MLLW)
or mean high water (MHW) or mean tide level (MTL)
are usually calculated by averaging over 19 years of data
(to eliminate the 18.6-year lunar nodal variations as well
as meteorologically caused noise). Such datums may
slowly change over the years due to a change in sea level
(due to climate change) or due to slow vertical land move-
ment (e.g., due to glacial rebound or sediment compac-
tion), the latter looking like a sea level change because
the water level gauge is mounted on the moving land.
All the datums slowly move up and down with the long-
term changes in relative sea level.
However, in shallow-water areas, there can also be
a tidally induced change in “mean sea level” due to
nonlinear tidal effects. In this case, what looks like
a “mean” result comes from averaging an asymmetry
within a tidal cycle caused by the nonlinear effects, rather
than from a uniform shifting up or down of the water level
on which the tide propagates. But this is an effect that is
still included in the datum calculation. It is therefore an
effect that should be considered when assessing long-term
changes in sea level in a shallow-water area, since some-
thing as simple as dredging or shoaling can change the
tidal hydrodynamics of a waterway and then also affect
the value of mean sea level (and thus all the tidal datums).
As one example, in the Delaware River and Bay, shallow-
water nonlinear effects increase the mean sea level value
near Philadelphia by an amount equivalent to 8 % of the
M2 amplitude at the entrance to the bay (Parker, 2007).
Tidal currents do not require datums, the zero current
speed serving that purpose (if there is no mean current
from a nontidal source). If there is a mean permanent cur-
rent, such as due to a mean river flow or a mean wind drift,
then the tidal current oscillates about that mean flow. Or,
as seen in Figure 8 (fourth example), the entire tidal cur-
rent ellipse can be shifted in the direction of the mean flow.
But here again, the mean current can also be affected by
 TIDAL HYDRODYNAMICS
the nonlinear tidal hydrodynamics causing an asymmetry
within the tidal cycle. This can be due not only to
shallow-water effects but even more importantly to
nonlinear lateral inertia effects (that do not need shallow
water). Such tidally induced residual current effects are
discussed below, as one of the many phenomena that
appear in tidal currents but not in the tide.
Special aspects of tidal currents (compared with
the tide)
The tide is a scalar quantity, with a one-dimensional
movement, that is, simply moving up or down. The tidal
current, however, is a vector quantity, representing the
horizontal water flow in two dimensions, and because it
is a vector quantity, its movements are more complicated
than the tide (but still very predictable because it has the
same astronomical forcing).
There are situations where the tidal current flow can
look approximately one dimensional, namely, when the
estuary or waterway is very narrow. In this case, a plot
of changing tidal current looks like a tide curve except
the sine curve is above and below a zero speed line. This
is called a reversing tidal current. In one tidal cycle, the
current goes from its maximum positive value
(maximum flood) through the zero current speed (slack
water) to the maximum negative value (maximum ebb)
and back again through slack water to maximum flood.
The slacks are usually referred to as slack before flood
(SBF) and slack before ebb (SBE). [Alternative names
for the above include flood strength, ebb strength, slack
flood begins (SFB), and slack ebb begins (SEB)].
Most tidal currents, however, are rotary tidal currents,
that is, the direction of flow rotates 360o (around the com-
pass) over one tidal cycle. This rotation is due to both the
Earth’s rotation and certain geographic conditions.
Because the tidal current speed varies as this rotation takes
place, the shape that the tip of the current vector traces out
can be an ellipse of various widths or even almost a circle
(in the open ocean) when the speed remains approximately
the same throughout the cycle (Figure 8). Within an estu-
ary that has a more complicated geography, for example,
those with islands and various channels, the shape can
even be more complicated (such as the shape shown in
the fourth plot in Figure 8). Because of the elliptical shape,
one can still use the terms flood current (for the tidal current
flow into and up an estuary) and ebb current (for the tidal
flow down and out of an estuary). However, there will not
be a true slack water, because for that brief time, the flow
will be cross-channel, so we use the terms minimum before
flood (MBF) and minimum before ebb (MBE).
Dramatic spatial variation in tidal
currents: vertically
The spatial variation of tidal currents, both horizontally
(i.e., geographically) and vertically in the water column,
is much more complex than the spatial variation of tides.
The tide, being the one-dimensional movement of the
697
water surface, does not have a variation with depth
(although the vertical tidal movement of different water
density layers does change considerably with depth).
However, tidal currents vary considerably from the sur-
face to the bottom. Bottom friction is a major cause of this
variation. The tidal current is slowest near the bottom and
faster nearer to (but usually not right at) the water surface.
Usually the tidal current will turn (from flooding to
ebbing, or vice versa) earlier near the bottom than near
the water surface, so there will be times when the tidal cur-
rent is going in the opposite direction at one depth than at
another depth. One example of the variation in tidal cur-
rent speed across a cross section of Chesapeake Bay is
shown in Figure 9. In this figure, one sees the current
flooding (i.e., flowing up the bay) in the deeper depths
and ebbing (i.e., flowing down the bay) near the water’s
surface (except on the west side, where it is still flooding).
There are also other effects that can make the tidal cur-
rent vary vertically. Tidal currents can be modified by
baroclinic effects, that is, by density differences vertically
along the water column, due to salinity differences
(in estuaries that are not well mixed) or temperature differ-
ences (offshore in the coastal ocean), which allow the
propagation of internal tide waves. Such baroclinic effects
on tidal currents are often seen in the middle of a stratified
water column, but not near the bottom or near the water
surface, where frictionally caused mixing takes place.
Dramatic spatial variation in tidal
currents: horizontally
Horizontally (i.e., geographically), the tide varies quite
smoothly due to the hydrodynamic effects of such things
as changing depths and widths, resonance, and Coriolis
force. Such variations can often be reproduced or
predicted with even simple analytical models. Although
tidal currents are also affected by these same hydrody-
namic effects and can change in similar ways, there are
other hydrodynamic effects which can make tidal currents
change dramatically in the horizontal direction, often over
surprisingly short distances. One example is the tidal cur-
rent in a navigation channel compared with the tidal cur-
rent in the nearby shallows. The tidal current is much
faster in the deeper channel than in the shallows, and the
times of slacks and of maximum floods and ebbs can be
quite different than those in the nearby shallows.
One does not require a dramatic change in depth to see
differences in the tidal currents. Even in a wide bay,
changes in bathymetry will affect the tidal currents. Not
only will current speeds vary with horizontal distance
(as mentioned above), but the bathymetry will steer the
current. Also, currents within a channel or constricted por-
tion of bathymetry will tend to be more reversing than cur-
rents in a more open and flat part of the bay, where a more
rotary tidal current will be possible.
In Figure 9, where vertical variation in the tidal currents
was seen, there was also horizontal variation along the
width of the bay. Bottom friction is often the main reason
for the horizontal variation in tidal current, due to the
698 TIDAL HYDRODYNAMICS
Tidal Hydrodynamics, Figure 9 Current flow at one moment in time through a cross section near the Chesapeake Bay Bridge.
The current is flooding up the Bay in the deeper depths and ebbing down the Bay near the water’s surface (except on the west side,
where it is still flooding).
changing water depth as one moves horizontally. How-
ever, there are many other causes of horizontal variation
in tidal currents, such as that discussed in the next section.
Nonlinear lateral inertial effects on tidal currents
One effect that can be even more dramatic than the fric-
tional effect (due to changing depths) is found where there
is a bending waterway, a channel bend, a point of land
projecting into the waterway, or some other similar geo-
graphic variation in the shoreline. Such a feature can cause
the formation of a large eddy during one or both phases of
the tidal current. A point projecting into a waterway pro-
duces a large eddy on the side of the point sheltered from
the tidal current. During the flood phase, there will be
a large eddy on the backside of the point, and during the
ebb phase, there will be a large eddy on the front side of
the point. This is a lateral inertial effect. After the ebb
phase ends, for example, the water on the backside of
the point keeps moving roughly in the same flood direc-
tion, because that location is sheltered from the opposing
flood currents by the point of land, and inertia keeps the
sheltered water moving (Parker, 2007).
If one harmonically analyzes current data from
a location within this eddy, one will obtain a consistent
mean current, which is usually referred to as a tidally
induced residual flow (and the process that causes it is
often called tidal rectification). However, one will also
see that the size of the tidal harmonic constants is very dif-
ferent than those for the waters not sheltered by the point
of land, because the inherent asymmetry leads to energy
being transferred to the second harmonics of the tidal con-
stituents. Thus, the ratio of semidiurnal-to-diurnal constit-
uents in the tidal current will be larger in sheltered
locations than in unsheltered locations (and thus, the typi-
cally shown diurnal-to-semidiurnal constituent ratio will
be smaller). Similarly the ratio of quarter-diurnal tidal cur-
rent constituents to semidiurnal tidal current constituents
will also be larger, leading to distorted tidal current curves.
None of this affects the tide, and one will not see such dra-
matic variations in tide constituents across the waterway.
This inertial effect is one of many nonlinear effects
which can modify tidal currents; however, this effect does
not depend on the water depth being shallow, as most
nonlinear effects do. When looking at the
two-dimensional or three-dimensional momentum equa-
tions (Parker, 2007), it is the lateral advective/inertial
terms that produce the tidally induced residual current
and the transfer of energy to higher frequency tidal constit-
uents. Figure 10 shows a good example of these inertial
effects on tidal currents, for the region where the Strait
of Juan de Fuca meets Haro Strait (between Canada and
the United States). Both are deep waterways, but they
meet at an angle, namely, the waterway bends signifi-
cantly to the north near Victoria, BC. Thus, the flood cur-
rent in the Strait of Juan de Fuca flowing past Victoria
cannot make a sharp left turn because inertia keeps it mov-
ing eastward, allowing the southerly ebb current at the
westernmost current station in Haro Strait to keep moving
longer than at the current station on the eastern side of the
waterway. Figure 10 shows harmonic analysis results
from three current stations across the entrance to
 TIDAL HYDRODYNAMICS 699

Tidal Hydrodynamics, Figure 10 Variation in (K1 + O1)/M2 and M4/M2 tidal current amplitude ratios, and in tidally induced residual
currents, at three stations across the entrance to Haro Strait at a depth of 70 ft (21 m) below MLLW.

Haro Strait (entering from the Strait of Juan de Fuca). The
(K1 + O1)/M2 amplitude ratio in the tidal current varies
dramatically, decreasing from 1.21 on the eastern side of
the waterway to 0.37 on the sheltered western side
(while for the tide the (K1 + O1)/M2 ratio does not change
much, only from 2.25 to 2.50). The M4/M2 ratio for the
tidal current also varies significantly across the waterway,
from 0.075 on the eastern side to 0.314 on the western
side.
Tidal whirlpools
The most dramatic horizontal variation in a tidal current
occurs in narrow straits (where the tidal current is very
fast) that suddenly widens out, reducing the flow
eventually but also producing a tidal whirlpool (also called
a maelstrom). This is a violently rotating funnel-shaped
hole of water that can suck ships underwater. To briefly
explain how the tidal whirlpool is generated, we first note
that when the water flows from the wide part of the strait
into the narrow part, the current follows the shoreline
and converges to flow more rapidly in the narrow part.
But when the tidal current reverses and flows from the nar-
row part of the strait into the wide part, the rapidly flowing
current cannot suddenly make the sharp left or right turn
that would spread the flow over the whole width of the
strait. The current’s inertia keeps its flow going approxi-
mately straight down the middle. This allows water near
the edges of the strait to continue moving in the same
700 TIDAL HYDRODYNAMICS
direction it had been flowing (also due to its inertia), until
the bending shoreline forces that edge flow to bend to the
middle and meet the flow down the middle, creating
a rotating whirlpool.
The most famous tidal whirlpool is the one that was
once violent and dangerous in the Strait of Messina
(between Sicily and the southern tip of the Italian main-
land), which Homer depicted in his Odyssey as the second
of two monsters, Scylla and Charybdis, faced by Ulysses.
Another dangerous tidal whirlpool is the Malström in the
narrow strait between two of the southern Lofoten Islands
off Norway, which was written about by both Jules Verne
and Edgar Allan Poe (Parker, 2012).
Nonlinear shallow-water effects on tidal currents
The shallow-water nonlinear processes that affect the tide
(as discussed earlier) also affect the tidal currents, and in
many situations those shallow-water effects are seen to
be more dramatic in the tidal current than in the tide. The
asymmetric tidal current can have (1) a shorter flood phase
with higher speeds and a longer ebb phase with slower
speeds (called flood dominance because of the higher
flood current speeds), (2) a shorter ebb phase with higher
speeds and a longer flood phase with slower speeds (ebb
dominance), or (3) equal flood and ebb phases, but where
one of the slacks can last for a couple of hours. Asymme-
try in the tidal current is important in the transport of sed-
iment and pollutants. The transport of coarse sediment
depends on the maximum speeds achieved and so might
be transported up an estuary with a flood-dominant situa-
tion. Fine sediment stays suspended except near slacks, so
the case with longer slacks before ebb might lead to depo-
sition at that time. (See Speer et al., 1991, for more discus-
sion on flood and ebb dominance.)
The distortion in tidal currents can be greatly enhanced
by a strictly linear superposition effect that, in fact, will
not similarly enhance the distortion in the tide. An extreme
example is in Ramshorn Creek, a small shallow channel
connecting the shallow Cooper and New Rivers, both part
of the Intracoastal Waterway in South Carolina and both
connected to the Atlantic Ocean (Parker, 1991). Two tide
waves, each distorted by shallow water, enter Ramshorn
Creek at opposite ends and cross, superimposing their
effects. The M2 flood currents for the two waves are in
opposite directions, and so the superposition of the two
waves leads to a reduction in the M2 tidal current. Each
tide wave is distorted to a different degree, because the
two waterways leading to Ramshorn Creek do not have
exactly the same width or depth. Thus, in each waterway
there is a different phase relationship between M2 and
M4, so that when the M4 tidal constituent waves are
superimposed the two waves add together (rather than
cancel each other out like the two M2 constituent waves),
thus increasing the M4 tidal current. The result is the very
unusual situation of a dominant quarter-diurnal tidal cur-
rent, that is, the tidal current actually changes directions
eight times a day instead of four (namely, there are four
tidal current cycles per day instead of two, each cycle
being 6.21 h long). There are other waterways in the salt
marsh estuaries of South Carolina and Georgia that also
exhibit the effects of two tide waves entering from oppo-
site ends and crossing each other. Most are not as extreme
as in Ramshorn Creek, but the result is still a very distorted
tidal current curve. (This superposition effect does not
lead to a dominant M4 in the tide because the two M2 tides
being scalars add positively at the crossover point, not
negatively as with the M2 vector tidal currents.)
Summary
Although it is astronomical forcing (the gravitational
effects of the moon and sun on the oceans) that makes
the tides so predictable, it is the hydrodynamics (i.e., phys-
ics of the water movement) that determines the size and
timing of the tides. When we speak of the tidal hydrody-
namics of estuaries, we are speaking of shallow-water
tides, which have much larger tidal ranges and faster tidal
currents than open-ocean tides. In shallow water, the
hydrodynamics also transfers tidal energy, through various
nonlinear processes, to new frequencies. These so-called
shallow-water tidal constituents include higher harmonics
of particular astronomical constituents (called overtides),
which distort the simple sinusoidal shape of the tide and
tidal current curves that are found in the open ocean, often
causing asymmetry in the tidal cycle in an estuary. These
same nonlinear processes also lead to interactions between
the tide and nontidal phenomena such as river discharge
and wind-induced changes (e.g., storm surges). In this
entry, we have explained how each of these mechanisms
work and the importance of tidal hydrodynamics in mov-
ing the water and affecting the processes in estuaries.
Bibliography
Parker, B. B., 1991. The relative importance of the various nonlinear
mechanisms in a wide range of tidal interactions. In Parker, B. B.
(ed.), Tidal Hydrodynamics. New York: John Wiley and Sons,
pp. 237–268.
Parker, B. B., 1998. The Coriolis effect: motion on a rotating planet.
Mariners Weather Log, 42(2), 1373. August 1998.
Parker, B. B., 2007. Tidal Analysis and Prediction. NOAA Special
Publication NOS CO-OPS 3, National Ocean Service, NOAA,
Department of Commerce, Silver Spring, MD.
Parker, B. B., 2012. The Power of the Sea. New York: Palgrave-
MacMillan.
Speer, P., Aubrey, D. G., and Friedrichs, C. T., 1991. Nonlinear
hydrodynamics of shallow tidal inlet/bay systems. In Parker,
B. B. (ed.), Tidal Hydrodynamics. New York: Wiley,
pp. 321–340.
Cross-references
Mean Sea Level
Tidal and Nontidal Oscillations
Tidal Asymmetry
Tidal Datum
Tidal Hydrodynamics
Tidal Ranges
Tides
 TIDAL AND NONTIDAL OSCILLATIONS
TIDAL AND NONTIDAL OSCILLATIONS
Bruce Parker
Center for Maritime Systems/Davidson Laboratory,
Stevens Institute of Technology, Hoboken, NJ, USA
Definition
Tidal oscillation is the oscillation in the vertical and hori-
zontal movement of water (and the properties of water)
caused by the tidal forces of the moon and sun.
Nontidal oscillation is the oscillation in the vertical and
horizontal movement of water (and the properties of
water) caused by nontidal phenomena that affect the estu-
ary and ocean, such as wind, atmospheric pressure, air
temperature, river runoff, and earthquakes.
Introduction
The movement of water into, out of, and inside an estuary
is critically important for understanding all processes that
take place in that estuary, whether they be physical, geo-
logical, chemical, or biological. There are numerous
examples of water movement effects on such processes,
only a few of which include transporting nutrients, flush-
ing pollutants, moving sediments, moving floating eco-
systems, changing the characteristics of water in which
an ecosystem occurs, eroding shorelines, mixing water
masses, and affecting stratification.
Oscillations
Many of the most important water motions involve some
type of oscillation, where there is movement around some
mean value (an equilibrium), that movement fluctuating
between positive and then negative values around that
mean. This can take many forms, such as the oscillation
of the water surface up and down (around a mean sea
level), water repeatedly flowing up and then down an estu-
ary, water temperature increasing and then decreasing,
salinity increasing and decreasing, as well as fluctuations
in numerous chemical, biological, and geological charac-
teristics affected by these physical oscillations. Properties
in an estuary undergo oscillations caused by a number of
the physical phenomena mentioned below.
Many of the most important of these oscillations are the
result of some type of wave motion. A wave is essentially
an oscillation that travels through space (in this case
through water or along the water surface), transferring
energy from one location to another and often transporting
mass (there are mass oscillations, but with the nonlinear
systems inherent in shallow estuaries, there is usually
overall transport of mass and other properties after each
complete oscillation). These waves can be of different fre-
quencies (periods) and wavelengths, with different driv-
ing/restoring forces. Some of these waves (e.g., the long
tide wave generated by the moon and sun in the open
ocean) are forced waves, where their generation is contin-
uous. Most other waves are free waves, where an initial
generating force sets them in motion, to either travel as
701
far as they can or cause oscillations in an estuary as long
as they can, using that initial input of energy.
Long waves
The most powerful of these waves tend to be long waves,
that is, waves whose wavelength (the distance from one
peak to the next peak) is much greater than the water depth
of the estuary (or for some long waves, even much greater
than the depth of the ocean to which the estuary is
connected). Most important are the tides, storm surges,
nearshore wind waves, and tsunamis (although the wave-
lengths of nearshore wind waves are not nearly as long
as for the other three types). Each of these long waves
has a different generating force.
In their largest form, each of these long waves can be
the cause of deadly natural disasters along the coast and
inside estuaries. Although the easy and accurate prediction
of tides makes tidal extremes less dangerous today, in the
past large tides, tidal bores, and tidal whirlpools killed
many people. Storm surges have killed many millions
over the centuries, because they were not predictable and
people living in low-lying coastal areas drown in the
floods caused by the storm surges. That did not change
until late in the twentieth century when satellites and
hydrodynamic computer models allowed us to accurately
predict where and when a storm surge would strike
and then provide warnings to those in the danger area.
Tsunamis are still not predictable (because the submarine
earthquakes that cause them are not predictable), which
is why we lost 300,000 people to a tsunami as recently
as 2004 in countries around the Indian Ocean or 25,000
people even in a tsunami-knowledgeable country like
Japan in 2011. Because of much better weather prediction,
we can now predict when large wind waves are likely to
strike the coast. For a history of marine disasters
and a history of how scientists slowly developed
a marine prediction capability, see Parker (2012).
In their less extreme form, tides, storm surges, and near-
shore wind waves have very significant day-to-day effects
on an estuary. They account for most of the water move-
ment in the estuary (and its effect on biological and geo-
logical systems) along with river discharge (which could
be thought of as having an annual asymmetric oscillation).
Many of these long wave phenomena can happen at the
same time, combining their effects. They also interact with
each other because the shallow water causes nonlinearity
in the motion. They do not just add to each other, they
change each other, by transferring energy from one to
another through various nonlinear mechanisms. (see Tidal
Hydrodynamics, and also Parker, 2007, for explanations
of mechanisms that cause the tide to interact with storm
surges, wind waves, and river discharge).
Types of oscillations
In most estuaries, the tides provide the dominant forces
that cause the water to move. The gravitational effects of
the moon and sun on the oceans cause the water to move
702 TIDAL AND NONTIDAL OSCILLATIONS
vertically (the tide) and horizontally (the tidal current), the
entire phenomenon usually referred to as the tides. Tidal
motion is actually the motion of extremely long waves
generated in the open ocean, which propagate over the
continental shelf and up into estuaries where their size is
usually amplified. Only the oceans are large enough for
the tide-generating forces to directly produce a tide of
significant size. The tides in an estuary are forced at the
estuary entrance by the tide wave from the ocean
(having been modified to some degree by propagating
over the shallower continental shelf). In estuaries we are
essentially speaking of shallow-water tides, which are sig-
nificantly different than open-ocean tides, having not only
much larger tidal ranges and faster tidal currents but also
distorted and asymmetric oscillations caused by the
nonlinear effects of the shallow water, which can have sig-
nificant effects on transport within the estuary (see Tidal
Hydrodynamics and also Parker, 2007).
Storm surges are very long waves, generated by the fast
winds of hurricanes and gales, which can push huge quan-
tities of water onshore causing extensive flooding. Such
wind-generated changes in water level along the shore
also occur under more modest wind conditions (under
conditions not considered as “storm” conditions), but the
term storm surge is often used to define in general any
change in water level caused by wind. Atmospheric
pressure also plays a small role in storm-induced changes
in water level. The crest of a storm surge does the most
damage when arriving at the coast at high tide. Because
of the shallow water, storm surges can modify the tide,
sometimes decreasing the tide range when the storm surge
has made the water depth greater (Parker, 2007).
Nearshore wind waves are also generated by wind, some-
times local winds, and sometimes by distant storms whose
waves travel across the ocean, reaching distant shores as
swell, which are deep-water waves until they reach the shal-
low waters near the coast and inside the estuary. At this point
the bathymetry can focus wave energy via wave refraction,
increasing their size at shallow points along the shore, creat-
ing longshore currents and rip currents, eroding shorelines,
and mixing the water column. During storms, they do the
most damage when arriving at the shore at high tide. Wind
waves can also affect the tide by modifying the bottom fric-
tion that affects the propagation of the tide wave.
Tsunamis are very long waves generated most fre-
quently by submarine earthquakes (ones that have signifi-
cant vertical movement of the sea bottom) but also
occasionally by volcanic eruptions or submarine land-
slides. Their wavelengths are very long in the deep ocean
(hundreds of kilometers), where they also travel at great
speeds (e.g., 600 km/h). When tsunamis reach the
shallower continental shelf, their wavelength shortens,
their speed decreases, their height increases, and they send
large waves against the shore with periods usually on the
order of 20–40 min. They do the most damage when
arriving at the shore at high tide (Parker, 2012).
Beyond these four primary long-wave phenomena,
there are other phenomena that can cause oscillations
in an estuary. Examples are seiches, land breeze-sea
breeze, and river flow.
Seiches are oscillations in harbors or small bays caused
by wind waves at the harbor entrance. These oscillations
are largest if the period of the waves entering the harbor
matches the natural period of the harbor basin
(as determined by its depth and width).
Land breeze-sea breeze oscillations in the wind are
found in tropical regions and in temperate regions during
the summer. During the day, the land heats up more than
the sea and the air rises above the land to be replaced by
cooler air blowing toward the land from the sea. The oppo-
site breeze occurs at night when the sea is warmer than the
land. A land breeze-sea breeze, which has roughly a 24-h
period, can produce an oscillating wind current in the
upper waters of the estuary or along the coast or even an
oscillation in the water level (although these oscillations
can be hidden by diurnal tidal oscillations). There are also
changes in wind speed and direction as weather systems
move over an estuary that can be quasiperiodic and can
cause oscillations in the estuary.
The daily change in air temperature also produces
a 24-h oscillation in the heating of the water surface,
which affects the estuary as does the 24-h cycle in the
effect of light on photosynthesis in phytoplankton. There
is, of course, an annual (seasonal) oscillation in tempera-
ture (and other meteorological effects).
River flow can be thought of as having a very asymmet-
ric oscillation, with most high flow values occurring after
high spring runoff from melting snow or from rainfall pro-
duced by large storms or hurricanes, with the remaining
time being characterized by much less flow. River currents
augment the tidal current, making the ebb flow duration
much longer and the flood duration much shorter.
The river flow can also interact with the tide nonlinearly,
reducing the tide range and distorting the shape of the tide
curve (see Tidal Hydrodynamics and Parker, 2012).
Summary
Tidal oscillations are oscillations in the vertical and hori-
zontal movement of water (and the properties of water)
caused by the tidal forces of the moon and sun. Nontidal
oscillations are horizontal and vertical oscillations in
water and its properties caused by nontidal phenomena
that affect the estuary and ocean, such as wind, atmo-
spheric pressure, air temperature, river runoff, and earth-
quakes. This entry summarizes the effects of tides, storm
surge, long wind waves, and tsunamis (all of which have
long wavelengths), as well as other phenomena that can
cause oscillations in an estuary, including seiches, land
breeze-sea breeze, daily and annual changes in air temper-
ature and light, and river discharge. Various aspects of the
biological, chemical, geological, and physical environ-
ments of an estuary have equilibria around which there
are oscillations caused by the above phenomena. Exam-
ples of these oscillations can be found elsewhere in this
encyclopedia.
 TIDAL RANGES 703

Bibliography Differences in tidal range are important, as they are often

Parker, B., 1998. The perfect storm surge. Mariners Weather Log, related to variations in coastal processes and morphology
44(2), 4–12. (Davies, 1980).
Parker, B., 2007. Tidal Analysis and Prediction. NOAA Special
Publication NOS CO-OPS 3, National Ocean Service, NOAA,
Department of Commerce, Silver Spring, MD. Description
Parker, B., 2012. The Power of the Sea. New York: Palgrave-
MacMillan. Temporally, tidal range varies due to changes in the posi-
tion and alignment of the moon and sun relative to the
earth. Every fortnight, the largest tidal ranges in semidi-
Cross-references urnal regions occur during spring tides, when the moon
Mean Sea Level and sun are in phase around times of new or full moon;
Seiche while the smallest tidal ranges occur during neap tides,
Storm Surges when the moon and sun are out of phase. In diurnal
Tidal Hydrodynamics
Tides regions, the largest tidal ranges occur every fortnight
during equatorial tides, when the moon is over the tro-
pics; while the smallest tidal ranges occur during tropic
tides, when the moon is over the equator. Over a month,
tidal range changes as the moon moves from its closest
TIDAL RANGES (perigee) approach to the earth to its furthest approach
(apogee) and back. Over annual time scales, changes in
Ivan D. Haigh tidal range occur as the sun’s position varies north or
Ocean and Earth Science, National Oceanography Centre, south of the equator and as it moves from its closest
University of Southampton, European Way, (perihelion) to furthest approach (aphelion) to the earth
Southampton, UK and back. The largest semidiurnal tidal range occurs in
March and September during the equinoxes, while the
Definition largest diurnal tidal range occurs in June and December
Tidal range is the vertical difference in height between during the solstices. Over longer time scales, variations
consecutive high and low waters over a tidal cycle. The in tidal range arise as a result of the 8.85-year cycle of
range of the tide varies between locations and also varies lunar perigee and the 18.61-year lunar nodal cycle
over a range of time scales (Stembridge, 1982). (Haigh et al., 2011).

Tidal Ranges, Figure 1 Global distribution of mean tidal range (Estimated using tidal constituents derived from TPXO7.2 global
ocean model (Egbert and Erofeeva, 2002) and downloaded from http://volkov.oce.orst.edu/tides/).
704 TIDAL REFLECTION
Spatially, tidal range varies according to the hydrody-
namic response of a particular ocean basin, shelf sea,
bay, or estuary to astronomical tidal forcing (Figure 1).
Tidal ranges are typically smallest in the open ocean,
along open ocean coastlines, and in almost fully enclosed
seas, such as the Mediterranean. Conversely, tidal ranges
are usually largest in semi-enclosed seas and funnel-
shaped entrances of bays and estuaries, such as the Bay
of Fundy in Canada or Bristol Channel in the UK, or
regions where a continental shelf has the right combina-
tion of depth and width for tidal resonance to occur, such
as on the northwest Australian shelf. Tides are often
crudely classified by their mean range into macrotidal
(>4 m), mesotidal (2–4 m), and microtidal (<2 m).
Bibliography
Davies, J. L., 1980. Geographical Variation in Coastal Develop-
ment. New York: Longman.
Egbert, G. D., and Erofeeva, S., 2002. Efficient inverse modeling of
barotropic ocean tides. Journal of Atmospheric and Oceanic
Technology, 19(2), 183–204.
Haigh, I. D., Eliot, M., and Pattiaratchi, C., 2011. Modeling global
influences of the 18.6-year nodal cycle and quasi-4.4 year cycle
on high tidal levels. Journal of Geophysical Research-Oceans,
116, C06025, doi:10.1029/2010JC006645.
Stembridge, J. S., 1982. Tidal range and variation. In Schwartz,
M. L. (ed.), Encyclopedia of Beaches and Coastal Environ-
ments. Dordrecht: Springer.
Cross-references
Tidal Datum
Tides
TIDAL REFLECTION
Manuel Díez-Minguito, Asunción Baquerizo Azofra,
Miguel Ortega-Sánchez and Miguel A. Losada Rodríguez
Andalusian Institute for Earth System Research,
University of Granada, Granada, Spain
Synonyms
Tidal wave reflection
Definition
Tidal reflection is a transformation process experienced by
a tidal wave in which all or part of its energy bounces back
due to geometric variations in the boundaries.
Description
Reflection is a common phenomenon experienced by
waves, and water waves are not an exception. Tidal wave
reflection occurs in a wide range of spatial scales, ranging
from tens of meters to thousands of kilometers. Semi-
enclosed bodies of water, such as estuaries, allow
(co-oscillating) tides to enter through their open bound-
aries. As the tidal wave propagates upstream, bathymetric
and topographic changes and the presence of obstacles
bounce back all or part of the incoming tidal energy. Prom-
inent tidal reflections typically occur at abrupt channel
constrictions and at upstream dams and weirs (e.g., in
the Guadalquivir estuary in Spain and in the Ems in the
Netherlands). Tidal reflection changes the structure of
tidal currents and ranges, which in turn affects the dynam-
ics of sediments and other solutes, and may also generate
residual movements with consequences in the long-term
morphology, water quality, and biota.
The prototypical case of tidal reflection is that of
a monochromatic tidal wave propagating into a straight,
frictionless channel of constant depth and width that is
closed at its head with an impermeable barrier. When the
traveling tidal wave encounters the head, all its energy is
reflected back as a sinusoidal wave of the same amplitude
and frequency that propagates in the opposite direction.
Physically, the oscillation resulting from their superposi-
tion is a standing wave with an antinode at the head, i.e.,
a position where the current vanishes and the amplitude
is twice the amplitude of the incoming wave. If the semi-
enclosed body is long enough in comparison to tidal
wavelength, at even/odd multiples of a quarter of the
wavelength off the head, antinodes/nodes are observed.
A tidal node is a location where the free surface does not
vary (null tidal elevation), and maximum along-channel
current amplitude occurs. Because the wave length
depends on the tidal period, different constituents will
exhibit different nodal point locations. For standing
waves, the tidal elevation and the current velocity are
p/2 out of phase. In fact, the character of the tidal wave
is usually analyzed (although not uniquely characterized)
by comparing the phase lag between the high water and
the high water slack with the values that would be
obtained under ideal conditions for standing waves
(0) and progressive waves (T/4), where T is the tidal
period. Resonance phenomena are also possible and not
infrequent in estuaries. For a given constituent, resonance
may occur if the estuary length L is about a quarter wave-
length of the forcing tide or odd multiples of thereof. This
means that the ratio of the maximum tide at the closed end
to the tide at the entrance becomes large. Perhaps the most
dramatic example of quarter-wave resonance is that
observed for the M2 semidiurnal tidal constituent in the
Bay of Fundy (Canada).
Friction dampens both the incident and reflected wave;
modifies their wave numbers, which depend on the along-
channel position; and also moves the position of nodes
and antinodes. If dissipation is significant and the estuary
is long enough, tidal energy may not reach the closed end
and no reflected wave is generated there. Even though the
incoming wave arrives with sufficient energy to the closed
end, the effective impact of tidal reflection may be limited
to a fraction of the estuary. The region of influence of tidal
reflection can be delimited by the position at which the
ratio between the amplitudes of the reflected and the inci-
dent waves is small. This practical approach requires sep-
arating the incident and the reflected wave by means of
 TIDAL REFLECTION 705

Tidal Reflection, Figure 1 Worldwide amphidromic systems for the semidiurnal M2 tide (12.42 h) obtained from Topex/Poseidon
satellite observations. Color contours indicate co-range areas in cm, and white lines are co-tidal lines. The amphidromic point east of
Chile coast shows the time of high water at each hour in the tidal cycle. Arrows show the clockwise or anticlockwise character of
several amphidromic points (Image adapted from http://svs.gsfc.nasa.gov/stories/topex/index.html by Richard Ray, Space Geodesy
branch, NASA/Goddard Space Flight Center).

analytical and observational methods. Tidal phases easily
obtained from harmonic analysis also provide useful infor-
mation because they tend to flatten out near the reflector
(different positions reach high water or low water at the
same time) when reflection dominates over friction. The
wave number obtained directly from the observed tidal
phases is thus typically much smaller than the wave num-
bers of the incident and reflected waves. Frictional effects
can significantly change the resonance condition and the
maximum amplification. Nevertheless, due to complex
bathymetry, and the variety of forcings involved, a precise
estimate often requires computational techniques.
In many natural situations, reflection may also intro-
duce a phase shift between the incoming and reflected
waves that affects the position of the antinodes and nodes.
Moreover, only a fraction of the incoming wave is
reflected. Energy reflection is characterized by
a (complex) reflection coefficient R, which is simply the
ratio between reflected and incident wave amplitudes.
Due to the imperfect reflection (R < 1), there are no true
nodes or antinodes in the wave profile, but quasi-
antinodes (maxima of the tidal elevation) and quasi-nodes
(minima of the tidal elevation). In simple terms, the
resulting wave is a partially (instead of fully) standing
wave, made up of a combination of a standing wave plus
an incoming progressive wave. In general, the reflection
coefficient is frequency dependent and varies spatially
with the absolute distance to the head due to the increase
in the dissipation and inverse shoaling of the reflected
wave. Nonlinear interactions between the two waves and
with other constituents may also be significant.
Tidal reflection also occurs due to topographic changes,
gradually as the wave propagates (e.g., variations in depth,
channel width, presence of meanders with small radius of
curvature, etc.). If the length scale of the spatial changes in
section is much larger than the tidal wavelength, energy
losses due to reflection are negligible and Green’s law,
which simply relates changes in channel width and depth
to changes in tidal range, is applicable. However, this
occurs rarely. In most cases tidal reflection is important
(e.g., at the sill of a fjord), and Green’s law is for practical
purposes inapplicable. Wave reflection from the margins
is also expected due to the geometry variations. For
instance, channel convergence causes partial reflection
of the incoming wave, thereby losing energy (apart from
bottom friction) as it propagates upstream. The seaward
radiation of (reflected) tidal energy due to channel conver-
gence and, if any, reflection at the upper limit may modify
the ocean tide near the basin entrance, especially when the
basin is close to resonance.
The basic tidal movement in the open ocean is also
a system of standing wave oscillations. Oceanic bound-
aries (mainly continents) reflect part of the tidal wave
energy, setting up nodal points, and also resonant motions,
if conditions exist for them. This reflection is responsible
for the differences of timing and tidal range along the
coasts, which in turn influence to a greater extent the estu-
ary mouth morphology (e.g., wave- or river-dominated
embayments occur where tidal action is weak). The
constraining effects of the emerged lands and the influence
of the Coriolis force result in the development of
amphidromic systems (Figure 1), in each of which the
tidal wave propagates around each tidal cycle. The range
of the rotating tide depends on the distance from the
amphidromic point: the larger the distance, the higher
the range. Co-range lines join places with the same tidal
range. These lines form more or less concentric circles
around an amphidromic point. Co-tidal lines, which link
all the points where the tide is at the same tidal phase,
radiate from the amphidromic point. Tidal waves in
706
amphidromic systems are typically Kelvin waves, namely,
gravity waves influenced by earth rotation. They propa-
gate along the coast, leaving it to the right (left) in the
northern (southern) hemisphere when facing in the direc-
tion of wave propagation. Their tidal ranges decrease
exponentially with increasing distance to the coast, and
currents are in geostrophic balance in the direction perpen-
dicular to the propagation. With a few exceptions, the tidal
waves of amphidromic systems tend to rotate anticlock-
wise in the northern hemisphere and clockwise in the
southern hemisphere. Again, local resonance is possible
if the dimensions of the basin match the tidal wavelength.
For instance, the North Sea provides a good example of
a resonant amphidromic system.
Summary
Tidal wave reflection is a quite common transformation
process experienced by a tidal wave and occurs simulta-
neously in a wide range of spatial scales. Sometimes
the reflection is prominent (in terms of fraction of
reflected energy), as for instance at a head weir, but some-
times is subtle, as occurs when tides propagate into
a long, weakly convergent estuary. This phenomenon
can even change the wave nature. Tidal reflection may
generate residual movements which in turn influence to
a greater extent long-term estuarine morphology and
water quality.
Bibliography
Officer, C. B., 1976. Physical Oceanography of Estuaries (and
Associated Coastal Waters). New York: Wiley.
Ray, R., 2007. NASA-GSFC, NASA-JPL, Scientific Visualization
Studio, and Television Production NASA-TV/GSFC. TOPEX/
Poseidon: Revealing Hidden Tidal Energy.
Valle-Levinson, A. (ed.), 2010. Contemporary Issues in Estuarine
Physics. Cambridge: Cambridge University Press.
TIDES
Ivan D. Haigh
Ocean and Earth Science, National Oceanography Centre,
University of Southampton, European Way,
Southampton, UK
Definition
Tides are the regular and predictable rise and fall of the sea
caused by the gravitational attraction and rotation of the
earth, moon, and sun system.
Introduction
Tides are the regular and predictable rise and fall of the sea
caused by astronomical forcing. The study of tides has
a long history and is perhaps the oldest branch of physical
oceanography (Cartwright, 1999). One of the fascinations
of tides is the number and diversity of the practical and
TIDES
scientific applications (Pugh, 1987). These include con-
cern with problems of navigation, coastal flooding and
erosion, transport of pollutants, and the extraction of tidal
power. Tides also have a controlling influence on geolog-
ical, sediment transport, water quality, and marine biolog-
ical processes.
Tides are normally used to refer to the vertical change
in sea level, whereas the term tidal currents is used for
the horizontal movement of water (Parker, 2005). Sea
level relates to the vertical change in the height of the
sea surface which occurs over all time and space scales
from many different mechanisms (including waves,
seiches, storm surges, tsunamis), with tides being the
most predictable and the dominant component of
sea-level variability in many parts of the world’s oceans
and coasts. Water level is the height of the sea surface
above some reference level or benchmark, often called
a tidal datum. Mean sea level is the average height of
the sea over longer periods of time (usually a month or
year), and hence the shorter-term variations of the tide
are filtered out.
Tide-generating force
Newton’s laws of gravitation attraction and motion form
the basis for physical understanding of how tides are gen-
erated. The universal law of gravitational attraction states
that the force (F) of attraction of two particles of masses
M1 and M2 separated by distance R is
M 1M 2
F¼G ð1Þ
R2
where G is the universal gravitational constant. Newton’s
laws of motion assert that a body (e.g., an element of sea-
water) remains at rest or constant velocity, unless acted
upon by a force, and that force is the product of a body’s
mass and acceleration.
To begin, consideration is given to the moon’s role in
generating tides. The earth and moon form a single sys-
tem, mutually revolving around a common center of mass
(known as the barycenter), with a period of 27.32 days.
The earth is 81 times larger than the moon, and as
a result the location of the barycenter lies within the earth.
The earth and moon circle about the barycenter, without
any rotation in absolute space. Therefore, each point on
the earth travels in circles which have the same radius.
Hence, they experience an equal centrifugal force (Fc),
directed parallel to a line joining the centers of the earth
and moon. While the centrifugal force is the same every-
where on the earth, the gravitational force varies with loca-
tion and is directed toward the center of the moon. The
local gravitation force (Fg) is given by
M eM m
Fg ¼ G ð2Þ
ðR  rÞ2
where Me and Mm are the mass of the earth and moon,
respectively, and r is the distance between the earth’s
 TIDES 707

a
20
Centifugal
15 Gravitational
Tide Generating
10

−5

−10

−15

−20
−40 −30 −20 −10 0 10 20 30 40 50 60
Centifugal Gravitational
Force Force
Gravitational Centifugal
Force Force
Centifugal Gravitational
Tide Generating Force Force Tide Generating
Force Force

Tide Generating
Force
b

Tides, Figure 1 (a) The magnitude and direction of the tide-generating force on the earth (looking side on to the earth), created by
the balance between the centrifugal force and the gravitational force. (b) The horizontal component of the tide-generating force
which causes water movement.

center and the point of interest. Locations nearer the moon
experience a local gravitational force that is larger than
that experienced at locations farther away.
For the earth-moon system to remain in equilibrium,
the total centrifugal force must exactly balance the force
of gravitation attraction, or else the earth and moon
would accelerate away or toward each other. At the center
of the earth the two forces balance, but at locations nearer
to the moon the gravitation force is larger than the centrif-
ugal force, and for locations farther away, the centrifugal
force is larger than the gravitational force (Figure 1a).
This difference is the tide-generating force (Ft), which
708
at a point on the earth is the difference between (1) and
(2), which simplifies to
M eM m
F t ¼ Ga ð3Þ
R3
where a is the radius of the earth.
The local variation in the tide-generating force does not
cause the water on the earth’s surface to be drawn into two
bulges on opposite sides of the earth. Instead it is the small
horizontal component of the force which causes water
movement (Figure 1b). This force, although small, has
nothing to oppose it; whereas the vertical component of
the tide-generating force acts against the much larger
gravitational attraction acting upon the ocean.
The solar tidal force can be thought of in a similar
manner. The mass of the sun is much greater than the mass
of the moon, but this is offset by the greater distance from
the earth. Therefore, the solar tidal forces are a factor of
0.46 weaker than the lunar forces.
Equilibrium tidal theory
To improve understanding of tides, Newton developed the
equilibrium tidal theory, in which three main assumptions
are made: (1) the earth is covered by an ocean of uniform
depth with no land masses; (2) the ocean responds imme-
diately to the tide-generating force; and (3) the effects of
rotation and friction can be ignored (Masselink et al.,
2003). Let us assume a stationary moon, aligned with the
earth’s equator. The lunar tide-generating force would
cause bulges on opposite sides of the earth in our uni-
formly deep ocean (Figure 2a). The earth rotates in an anti-
clockwise direction on its polar axis, beneath these bulges,
taking 24 h (a solar day) to complete one rotation. Hence,
anywhere on the earth’s surface you would experience two
high tides each day, but with decreased height as you
moved north or south of the equator (Figure 2b). In reality
by the time the earth has completed one full rotation, the
moon has moved on in its rotation around the barycenter.
It takes about 50 min longer than a solar day for the point
on the equator to reach the maximum part of the first bulge
again. This is why high semidiurnal tides occur every 12 h
and 25 min, not every 12 h.
At any point on the earth, the change in the height of the
tide can be described by a simple harmonic curve of the
form (Figure 2b, d)
H cos ðot  g Þ ð4Þ
where H is the amplitude, o is the frequency, g is the phase
lag relative to a defined time zero, and t is time. Assuming
that the moon rotates around the earth in a purely circular
path aligned with the equator, we term this harmonic M2.
If we were to ignore the moon and just consider the sun,
then the same features would be observed, except the time
between high waters would be exactly 12 h and the height
of high water would be just less than half of that observed
for the moon. This harmonic is termed S2.
TIDES
Now consider the combined effect of the moon and
sun. When the earth, moon, and sun are aligned during
full or new moon, the equilibrium tidal bulges associated
with the moon and sun are in phase and combine to
create a spring tide with a larger than average tidal range
(Figure 3). This occurs about every 15 days. When the
moon is positioned half way between the new and
full phases, the two tidal bulges are out of phase and
combine to produce a neap tide with smaller than average
tidal range.
Tidal patterns are complicated by the fact that for much
of the year the moon and sun are not aligned with the equa-
tor. The earth rotates around the sun on a plane called the
ecliptic, which is inclined at 23 270 to the equator. The
moon orbits the earth on a plane that is included by 5 90
to the ecliptic. As the moon revolves, its position above
the earth varies between latitudes 28 360 (23 270 + 5 90 )
north and south of the equator, and the tidal bulges are
titled relative to this angle (Figure 2c). An observer on
the earth’s surface would experience two high tides
a day, but the relative heights of each would be different,
with the relative differences increasing moving farther
north or south (Figure 2d). This so-called diurnal inequal-
ity reaches a maximum when the moon is at its maximum
northern or southern declination about every 2 weeks.
The sun’s declination varies over a 365.25-day cycle,
by 23 270 north or south of the equator, causing the sea-
sons. The largest diurnal tides occur in June and December
during the solstices and the smallest diurnal tides in March
and September during the equinoxes. As the strength of
the diurnal force increases, the semidiurnal force reduces,
and vice versa (Pugh, 2004). Hence, the semidiurnal tidal
forces reach their maximum at the equinoxes.
Tidal patterns are further complicated because the
orbits of the earth and moon are elliptical, not circular.
As a result the distance between the moon and sun with
the earth varies, altering the strength of the tide-generating
force. Over a period of 27.6 days, the moon moves
from perigee (its closest approach to the earth) to apogee
(its farthest approach to the earth) and back. At perigee
and apogee the lunar tidal forces are 15 % greater or less
than average, respectively. The orbit of the earth around
the sun is also elliptical, moving from perihelion (closest
to the sun) to aphelion (farthest from the sun) over
a period of 365.25 days. The difference in distance
between these two positions is only about 4 % so tides
are only marginally larger at perihelion compared to
aphelion.
Over longer time scales, variations in tidal forces arise
as a result of the 8.85-year cycle of lunar perigee (which
influences tides as a quasi 4.4-year cycle) and the 18.61-
year lunar nodal cycle (Haigh et al., 2011).
Harmonic analysis and tidal prediction
Predicting times and heights of tides has many practical
applications, and there has been a long history of develop-
ment from simple techniques that related the time of high
 TIDES 709

a b

3 3
2 2
1 1

2.5
c d

2 1

1.5
Level (m)

1 2

0.5

0 3

−0.5
0 0.5 1 1.5 2 2.5 3 0 0.5 1 1.5 2 2.5 3
Days Days

Tides, Figure 2 (a, b) The earth’s (looking side on to the earth) double tidal bulge aligned with the moon which is located over the
equator in (a) and over the Tropic of Cancer in (b). (c, d) Tidal curves (offset for presentation purposes) over a three-day period at sites
1, 2, and 3, the locations of which are marked on plots (a) and (b), respectively.

water to the phase of the moon (Pugh, 1987), through to the
more sophisticated harmonic (Doodson, 1921; Foreman,
1977) and response analysis methods (Munk and Cartwright,
1966). Harmonic analysis is the most widely used tidal pre-
diction method. The underlying principle of this method is
the assumption that no matter how complex they appear,
tidal variations at any location can be represented by the
sum of a finite number of harmonic terms (tidal constituents),
each expressed in the form of (4).
The frequency of each of the tidal constituents can be
expressed as a linear superposition of six fundamental
astronomical forcing harmonics, from which the positions
of the sun or moon, and hence the tide-generating forces,
can be calculated for any time. Therefore, it is possible
to relate each individual tidal harmonic to real astronomi-
cal behavior (Pugh, 2004). More than 400 tidal constitu-
ents have been identified (Cartwright and Tayler, 1971),
but the tide can be predicted to a useful level of accuracy
using only a small subset of these.
In order to predict tides at a given location, we need to
know the frequencies, amplitudes, and phases of the tidal
constituents. The frequencies are fixed, but the amplitudes
and phases of the different constituents vary with location
and the aim of the harmonic analysis is to determine them.
710 TIDES

a
Spring tides

b
Neap tides

c
3

2.5

1.5 Lunar Tide (M2)

Level (m)

1
Solar Tide (S2)
0.5
Neaps
0

−0.5
Springs Combined Tide (M2 + S2)

−1
0 5 10 15
Days

Tides, Figure 3 View looking down upon the earth from the north pole when (a) the earth, moon, and sun are aligned (during full
or new moon) – the equilibrium tidal bulges associated with the moon and sun are in phase and combine to create a spring tide;
(b) the moon is positioned half way between the new and full phases – the two tidal bulges are out of phase and combine to produce
a neap tide with smaller than average tidal range. (c) Lunar, solar, and combined tidal curves (offset for presentation purposes)
over a 15-day period at a point on the equator.

This involves least-squares fitting of the tidal constituents
to water level measurements at that site. To predict the
tides into the future, the estimated amplitudes and phases
of each constituent can be substituted into (4) and then
the terms summed.
Dynamic tidal theory
The equilibrium theory helps to describe several features
of tides and serves as a reference system for harmonic
analysis. However, the tide observed in reality bears no
resemblance to the equilibrium tide, because none of the
 TIDES
Tides, Figure 4 Global distribution of the tidal form factor (Estimated using tidal constituents derived from TPXO7.2 global ocean
model (Egbert and Erofeeva, 2002) and downloaded from http://volkov.oce.orst.edu/tides/).
principle assumptions of the theory are valid (Brown et al.,
1989), including the following: (1) the presence of land
prevents tidal bulges from directly circumnavigating the
globe; (2) the rotation of the earth on its polar axis is too
rapid for the inertia of the water masses to overcome in
sufficient time to establish an immediate equilibrium tide;
and (3) water movements induced by tide-generating
forces are subject to friction and the Coriolis force.
The combination of all of these factors results in the
development of so-called amphidromic systems. The
global ocean is essentially broken into separate basins sep-
arated by shallow continental shelves and landmasses
(Masselink et al., 2003). In these enclosed basins, the tidal
wave travels from east to west, elevating sea levels against
the western margin of the basin. The resulting slope in the
ocean surface produces a pressure gradient that causes
water to flow eastward. In the Northern Hemisphere, the
eastward flow is deflected to the right by Coriolis force,
which causes elevated sea levels against the southern
margin. The pressure gradient force subsequently drives
the water northward which is then deflected to the west
and so on. Thus, the wave crest rotates around the ocean
basin in an anticlockwise direction, which is referred to
as a Kelvin wave. Amphidromic systems are set up on
continental shelves and large bays, but these systems are
scaled down and are less symmetrical as the wave speeds
decrease in shallow water where there are significant
energy losses due to bottom friction (Pugh, 2004).
In the open ocean, tides are generated directly by the
tide-generating forces. These waves then spread onto the
711
surrounding shallow shelf seas where their characteristics
are altered through standing wave generation and local
resonances (Pugh, 2004). As the tidal wave progresses
from the deep ocean onto the shallow continental shelf,
the wave slows down. The wavelength is reduced and
the amplitude increases because the wave energy becomes
concentrated in a smaller area. As the wave approaches
even shallower water near the coast, it encounters the
irregular topography of the coastline and moves into estu-
aries and bays where it undergoes even larger distortions.
As the tide travels into a narrowing bay or estuary, the tidal
range increases again, because the same energy is being
forced through a smaller opening (Parker, 2005). In
extreme cases, this can result in the formation of a tidal
bore. In very shallow water, the wave crest will travel
faster than the trough, resulting in tidal asymmetry. Even-
tually, the tidal amplitude will be reduced by bottom fric-
tion as the wave progresses into very shallow water.
Tidal characteristics and types
At a particular location, tides are typically classified by
their range (see Tidal Ranges) and form. The form of the
tide can be estimated from the tidal form factor (F). This
is derived from the main diurnal and semidiurnal tidal con-
stituent amplitudes (H) using the following equation
(Pugh, 2004):
ðH K 1 þ H O2 Þ
F¼ ð5Þ
H M 2 þ H S2
712 TIMESCALE
It can be used to quantify whether a site experiences
(Figure 4) the following: two high and low tides each lunar
day, semidiurnal tidal form (F < 0.25); a single high and
low tide, diurnal tidal form (F > 3); or periods when
both types occur, mixed tidal form (mixed, mainly
semidiurnal, F ¼ 0.25 to 1.50; mixed, mainly diurnal,
F ¼ 1.50 to 3.00).
Summary
To fully understand tides and predict them, one must under-
stand both the astronomical forcing that gives rise to them
and also the response of the oceans, bays, and estuaries to
this forcing (Parker, 2005). Knowledge of the astronomical
forcing helps to describe several features of tides and
servers as a basis for the tide’s predictability. However, it
is the knowledge of the hydrodynamics of the tide in
a particular ocean basin, shelf sea, or estuary which pro-
vides understanding of the timing of high and low water,
the size of the tidal range, the form of the tide, and any com-
plex shallow water distortions of the tidal curve.
Bibliography
Brown, J., Colling, A., Park, D., Philips, J., Rothery, J., and Wright,
J., 1989. In Bearman, G. (ed.), Tides. Waves, Tides and Shallow-
Water Processes. Oxford: Butterworth Heinemann and Open
University.
Cartwright, D. E., 1999. Tides, a Scientific History. Cambridge:
University Press.
Cartwright, D. E., and Tayler, R. J., 1971. New computations of the
tide-generating potential. Geophysical Journal of the Royal
Astronomical Society, 23(1), 45–73.
Doodson, A. T., 1921. The harmonic development of the tide-
generating potential. Proceedings of the Royal Society of London
Series A, 100(704), 305–329.
Egbert, G. D., and Erofeeva, S., 2002. Efficient inverse modeling of
barotropic ocean tides. Journal of Atmospheric and Oceanic
Technology, 19(2), 183–204.
Foreman, M. G. G., 1977. Manual for Tidal Heights Analysis and
Prediction. Pacific Marine Science Report No. 77–10, Institute
of Ocean Sciences, Patricia Bay, Sidney, BC. http://www.omg.
unb.ca/GGE/5013_LABS/heights.pdf.
Haigh, I. D., Eliot, M., and Pattiaratchi, C., 2011. Modeling global
influences of the 18.6-year nodal cycle and quasi-4.4 year cycle
on high tidal levels. Journal of Geophysical Research-Oceans,
116, C06025, doi:10.1029/2010JC006645.
Masselink, G., Hughes, M., and Knight, J., 2003. Introduction to
Coastal Processes and Geomorphology. London: Hodder Education.
Munk, W. H., and Cartwright, D. E., 1966. Tidal spectroscopy and
prediction. Philosophical Transactions of the Royal Society of
London, A259, 533–581.
Parker, B., 2005. Tides. In Schwartz, M. L. (ed.), Encyclopedia of
Coast Science. Dordrecht: Springer.
Pugh, D. T., 1987. Tides, Surges and Mean Sea Level: A Handbook
for Engineers and Scientists. Chichester: Wiley.
Pugh, D. T., 2004. Changing Sea Levels: Effects of Tides, Weather
and Climate. Cambridge: Cambridge University Press.
Cross-references
Seiche
Storm Surges
Tidal and Nontidal Oscillations
Tidal Asymmetry
Tidal Datum
Tidal Hydrodynamics
Tidal Ranges
TIMESCALE
Lisa V. Lucas
United States Geological Survey, Menlo Park, CA, USA
Definition
Time scale refers to “the time allowed for or taken by
a process or sequence of events” (OUP, 2013).
Essential concepts, application and usage
The above definition provides a general description of the
term “time scale.” In science and engineering practice, a
time scale is typically an estimate expressing a representa-
tive, overall magnitude as opposed to a precise value; as
such, time scales (like length, velocity, and other commonly
used scales) are frequently quantified in terms of orders of
magnitude (Cushman-Roisin and Beckers, 2011). A time
scale carries the dimension of time and may be expressed
either in precise temporal units (e.g., seconds, hours, days,
years, decades) or in approximate temporal units (as with
“tidal,” “seasonal,” or “episodic” time scales).
In combination with other scales, time scales may be
used to estimate the relative magnitude (and thus, impor-
tance) of individual terms in time-marching equations
(Cushman-Roisin and Beckers, 2011). Time scales may
be defined to describe the rates of physical, biological,
and chemical processes, serving as a common currency
with which to compare their relative speed, and thus
importance (e.g., Koseff et al., 1993; Middleburg and
Nieuwenhuize, 2000; Lucas et al., 2009; Dame, 2012).
In such a context, a smaller time scale suggests a faster
and more dominant process (Lucas, 2010). A time scale
may be estimated as the reciprocal of a first order rate con-
stant or frequency, or from a combination of length, veloc-
ity, diffusivity, or other scales such that the remaining
dimension is time (e.g., Fischer et al., 1979; Koseff et al.,
1993). A time scale may reflect the approximate time for
completion of a process, such as for (1) diffusive mixing
over an estuary’s depth (Fischer et al., 1979), (2) advective
transport over the estuary length (MacCready and Banas,
2011), (3) flushing of an estuary by river flow and/or tides
(Sheldon and Alber, 2006), (4) settling of particles through
a water column (de Brauwere and Deleersnijder, 2010),
(5) biomass growth by a factor e (Lucas, 2010), or (6) filtra-
tion of a water column or estuary volume by benthic
organisms (Buzzelli et al., 2013).
The term “time scale” may also be used to convey (1) a
typical period of variability or fluctuation in estuarine forc-
ing or ecosystem response (e.g., Litaker et al., 1993; Cloern,
1996; Jay et al., 2000), (2) a period of estuary adjustment to
 TOMBOLO
low-frequency forcing (e.g., Monismith et al., 2002;
MacCready and Geyer, 2010), (3) the period(s) of variabil-
ity captured by measurements or models (e.g., Blumberg
et al., 1999; Wang et al., 1999), or (4) the temporal lens
through which processes are examined (e.g., Stacey et al.,
2001; Chapin III et al., 2006).
Summary
Described very generally, a “time scale” is the time taken
by a process or sequence of events (OUP, 2013). However,
there exist many shades of meaning and usage of this term,
which is ubiquitous in engineering and increasingly so in
environmental science. Typically, a “time scale” implies
a time estimate, not a precise value. Comparison of time
scales for two or more coincident processes (be they phys-
ical, biological, or chemical) provides a diagnostic tool for
assessing the processes’ relative importance and, poten-
tially, for simplifying complex equations.
Bibliography
Blumberg, A. F., Khan, L. A., and St. John, J. P., 1999. Three-
dimensional hydrodynamic model of New York Harbor region.
Journal of Hydraulic Engineering, 125, 799–816.
Buzzelli, C., Parker, M., Geiger, S., Wan, Y., Doering, P., and
Haunert, D., 2013. Predicting system-scale impacts of oyster
clearance on phytoplankton productivity in a small subtropical
estuary. Environmental Modeling and Assessment, 18, 185–198.
Chapin, F. S., III, Woodwell, G. M., Randerson, J. T., Rastetter,
E. B., Lovett, G. M., Baldocchi, D. D., Clark, D. A., Harmon,
M. E., Schimel, D. S., Valentini, R., Wirth, C., Aber, J. D., Cole,
J. J., Goulden, M. L., Harden, J. W., Heimann, M., Howarth,
R. W., Matson, P. A., McGuire, A. D., Melillo, J. M., Mooney,
H. A., Neff, J. C., Houghton, R. A., Pace, M. L., Ryan, M. G.,
Running, S. W., Sala, O. E., Schlesinger, W. H., and Schulze,
E. D., 2006. Reconciling carbon-cycle concepts, terminology,
and methods. Ecosystems, 9, 1041–1050.
Cloern, J. E., 1996. Phytoplankton bloom dynamics in coastal eco-
systems: a review with some general lessons from sustained
investigation of San Francisco Bay, California. Reviews of Geo-
physics, 34(2), 127–168.
Cushman-Roisin, B., and Beckers, J.-M., 2011. Introduction to
Geophysical Fluid Dynamics: Physical and Numerical Aspects.
Waltham: Academic.
Dame, R. F., 2012. Ecology of Marine Bivalves: An Ecosystem
Approach, 2nd edn. Boca Raton: CRC Press.
de Brauwere, A., and Deleersnijder, E., 2010. Assessing the
parameterisation of the settling flux in a depth-integrated model
of the fate of decaying and sinking particles, with application to
fecal bacteria in the Scheldt Estuary. Environmental Fluid
Mechanics, 10, 157–175.
Fischer, H. B., List, E. J., Koh, R. C. Y., Imberger, J., and Brooks,
N. H., 1979. Mixing in Inland and Coastal Waters. San Diego:
Academic.
Jay, D. A., Geyer, W. R., and Montgomery, D. R., 2000. An ecolog-
ical perspective on estuarine classification. In Hobbie, J. E. (ed.),
Estuarine Science: A Synthetic Approach to Research and Prac-
tice. Washington: Island Press, pp. 149–176.
Koseff, J. R., Holen, J. K., Monismith, S. G., and Cloern, J. E.,
1993. Coupled effects of vertical mixing and benthic grazing
on phytoplankton populations in shallow, turbid estuaries. Jour-
nal of Marine Research, 51, 843–868.
Litaker, W., Duke, C. S., Kenney, B. E., and Ramus, J., 1993. Short-
term environmental variability and phytoplankton abundance in
713
a shallow tidal estuary. II. Spring and fall. Marine Ecology Pro-
gress Series, 94, 141–154.
Lucas, L. V., 2010. Implications of estuarine transport for water
quality. In Valle-Levinson, A. (ed.), Contemporary Issues in
Estuarine Physics. Cambridge: Cambridge University Press,
pp. 272–306.
Lucas, L. V., Thompson, J. K., and Brown, L. R., 2009. Why are
diverse relationships observed between phytoplankton biomass
and transport time? Limnology and Oceanography, 54(1),
381–390.
MacCready, P., and Banas, N. S., 2011. Residual circulation,
mixing, and dispersion. In Wolanski, E., and McLusky, D. S.
(eds.), Treatise on Estuarine and Coastal Science. Waltham:
Academic, Vol. 2, pp. 75–89.
MacCready, P., and Geyer, W. R., 2010. Advances in estuarine
physics. Annual Review of Marine Science, 2, 35–58.
Middleburg, J. J., and Nieuwenhuize, J., 2000. Uptake of dissolved
inorganic nitrogen in turbid, tidal estuaries. Marine Ecology Pro-
gress Series, 192, 79–88.
Monismith, S. G., Kimmerer, W., Burau, J. R., and Stacey, M. T.,
2002. Structure and flow-induced variability of the subtidal
salinity field in Northern San Francisco Bay. Journal of Physical
Oceanography, 32, 3003–3019.
OUP, 2013. Oxford Dictionaries. Oxford: Oxford University Press.
http://oxforddictionaries.com.
Sheldon, J. E., and Alber, M., 2006. The calculation of estuarine
turnover times using freshwater fraction and tidal prism models:
a critical evaluation. Estuaries and Coasts, 29(1), 133–146.
Stacey, M. T., Burau, J. R., and Monismith, S. G., 2001. Creation of
residual flows in a partially stratified estuary. Journal of Geo-
physical Research, 106(C8), 17013–17037.
Wang, L., Sarnthein, M., Erlenkeuser, H., Grimalt, J., Grootes, P.,
Heilig, S., Ivanova, E., Kienast, M., Pelejero, C., and Pflaumann,
U., 1999. East Asian monsoon climate during the Late Pleisto-
cene: high-resolution sediment records from the South China
Sea. Marine Geology, 156, 245–284.
Cross-references
Residence Time
TOMBOLO
Michel Michaelovitch de Mahiques
Oceanographic Institute of the University of São Paulo,
Sao Paulo, Brazil
Synonyms
Ayr (Northern Europe); Tie bar
Definition
A tombolo is a sediment deposit at the coast formed by
wave refraction and diffraction at the edges of an obstacle
(natural or artificial) originally detached from the
mainland.
Description
The formation of a tombolo is caused by the diffraction of
the wave at the ends of an obstacle, creating a convergence
of opposing flows of sediment transport. Depending on
714 TOXIC BLOOMS
the input of sediments and sea-level trends, the accumula-
tion of sediments (sands or gravels) leads to continuous
progradation of the coastline towards the detached
obstacle (usually an island). A smaller sediment protru-
sion can also be developed in the back side of the obstacle.
The final phase of this process is the complete coalescence
between these sediment features and the blockage of the
water body located behind the obstacle, with the conse-
quent formation of a connection between the obstacle
and the mainland (i.e., the tombolo). Tombolos can be
classified as simple, when the feature is formed by
a single sediment body, or complex (doubles, triples,
etc.), in which the sediment ridges can be formed in
a different time (Johnson and Reed, 1910). In this last
case, lagoons can be formed between ridges, marking
intervals of interruption of the tombolo formation.
Bibliography
Johnson, D. W., and Reed, W. G., Jr., 1910. The form of Nantasket
beach. Journal of Geology, 18, 162–189.
Cross-references
Coastal Lagoons
Coastal Landforms
Shoreline
TOXIC BLOOMS
Kevin G. Sellner
Chesapeake Research Consortium, Edgewater, MD, USA
Synonyms
Allelopathic algae; Harmful algal blooms (HABs);
High-biomass blooms; “Red tides”
Definition
Toxic blooms are the accumulation of toxin-producing
cells that lead to aquatic living resource mortalities or ill-
ness or deaths in avian or terrestrial mammals, including
humans.
Mortalities from blooms can also occur through indirect
effects of the accumulation of high biomass typical of
blooms and the subsequent respiration or death and
decomposition that consumes the available oxygen, lead-
ing to suffocation and dying and dead fauna.
Introduction
Toxin-producing taxa and syndromes
Toxic blooms are generally accumulations of phytoplank-
ton (microscopic single-cell free-floating plants) and
occasionally mixotrophic or heterotrophic dinoflagellates,
benthic dinoflagellates, or macroalgae. Some of the cells
accumulating produce complex intracellular compounds
that are toxic to various freshwater and marine fauna as
well as higher organisms that include birds and
mammals. The suite of toxins produced (http://www.
issha.org/Welcome-to-ISSHA/Harmful-Algae-Links) has
a primary toxin and then many substituted congeners
and include paralytic shellfish poisons (PSP), primarily
from saxitoxin and gonyautoxins and their derivatives in
marine dinoflagellates like Alexandrium species and
others; neurotoxic shellfish poisons (NSP) such as
brevetoxin produced by Karenia brevis in Florida red tides
and anatoxin from some freshwater cyanobacteria;
domoic acid for amnesic shellfish poisons (ASP) pro-
duced by some marine members of the diatom genus
Pseudo-nitzschia; ciquatera fish poisons (CFP) from toxin
precursors for ciguatoxins and maitotoxins produced by
some benthic marine dinoflagellate species like
Gambierdiscus; diarrhetic shellfish poisons (DSP) from
okadaic acid produced by the marine dinoflagellate
Dinophysis and ichthyotoxic karlotoxins from the brack-
ish dinoflagellate Karlodinium, which kill many fish and
dramatically reduce grazing pressures from small plank-
tonic grazers; azaspiracids which induce vomiting and
diarrhea, produced from the marine dinoflagellate
Azadinium spinosum; and the hepatotoxic microcystins
produced by Microcystis and other freshwater
cyanobacteria. Other toxins (prymnesins, nodularin,
yessotoxin, palytoxins, tetrodotoxin, spirolides) are also
found less frequently throughout the world. Exposure for
most birds and mammals is primarily through shellfish
and other organisms that concentrate toxic cells and are
subsequently ingested by the larger consumers, including
humans; drinking water supplies may also be toxin
sources for some of the freshwater toxins. Distributions
of these toxins and bloom formers can be viewed at
http://www.whoi.edu/redtide/regions/world-distribution.
High-biomass blooms
Toxin- and nontoxin-producing populations of the plank-
ton and macroalgae noted above can accumulate to such
high levels that the oxygen demand associated with the
very abundant organisms can consume all available
oxygen in surrounding waters, posing serious threats to
aerobic populations of fish, shellfish, and many benthic
taxa. These high-biomass blooms are increasingly com-
mon to estuaries, and shallow coastal lagoons receiving
high nutrient loads (see Anderson et al., 2002; Heisler
et al., 2008) and bloom formers include Microcystis and
Anabaena, Prorocentrum, Ceratium, Cochlodinium
polykrikoides, raphidophytes, pelagophytes (brown tides),
and other HAB species. Dense beds or scoured
macroalgae can also occur, leading to high respiratory
demand during decomposition and oxygen-poor condi-
tions that threaten coexisting fauna. Finally, spring diatom
blooms can be very large in temperate estuaries where
post-bloom deposition and decomposition lead to hypoxia
or anoxia in bottom waters (e.g., Chesapeake Bay, Kemp
and Boynthon, 1992); Asian countries typically include
diatoms in lists of HABs from regional waters.
 TOXIC BLOOMS
Toxicity from other compounds (allelopathy)
Many of the bloom-forming species produce other com-
pounds that are toxic to other members of the plankton,
whether other phytoplankton taxa that compete for nutri-
ents and sunlight in active growth or potential grazers that
could limit accumulation. These compounds are “toxic” in
that they provide the bloom former greater opportunity for
maintaining growth, while their competitors and predators
are limited in their ability to lower their growth rates or
accumulation. Referred to as allelopathic compounds,
several bloom taxa produce spectrums of compounds with
these properties. Examples include Karenia brevis, the
Florida red tide taxon, which produces compounds that
kill two competing diatoms and a dinoflagellate (Prince
et al., 2008) and retard rotifer grazing on the dinoflagellate
(Kubanek et al., 2007). Alexandrium tamarense and
A. minutum produce compounds that had negative effects
on the diatom Thalassiosira weissflogii and the
cryptophyte Rhodomonas, as well as a natural community
and a ciliate (Fistarol et al., 2004). Cyanobacteria also pro-
duce inhibitory compounds, with Nodularia spumigena,
Aphanizomenon flos-aquae, and Anabaena lemmermannii
all inhibiting these same taxa (Suikkanen et al., 2004).
Another dinoflagellate, Cochlodinium polykrikoides,
inhibits the growth of Akashiwo sanguinea, Gymnodinium
instriatum, and Chattonella marina as well as members of
the natural phytoplankton community from Long Island
estuaries likely through production of reactive oxygen
species (Tang and Gobler, 2010); raphidophytes like
C. marina, C. antiqua, Fibrocapsa japonica, and
Heterosigma akashiwo have similar ROS production
capacities (Marshall et al., 2005). ROS production can
lead to gill damage in fish and death (Kim et al., 1999).
The prymnesiophyte Prymnesium parvum produces com-
pounds inhibitory to several dinoflagellates (Fistarol et al.,
2003). Some macroalgae (Ulva, Corallina, Sargassum,
Gracilaria, many coral reef cyanobacteria) also produce
allelopathic compounds inhibitory to other phytoplankton
and macroalgae (Nagle and Paul, 1999; Wang et al., 2007;
Tang and Gobler, 2011).
Mechanisms for bloom formation
Planktonic blooms can originate from physical mecha-
nisms (Sellner et al., 2003) like circulation patterns
(Langmuir cells, fronts, upwelling), local storms and
precipitation, poor mixing, stratification, limited flushing,
or periods of low nutrient but abundant light which favors
surface accumulations via positive phototactic swimming
or buoyancy control of several toxic species. Examples of
these physically driven aggregations of cells include fron-
tal concentration of Gymnodinium in the Potomac River
estuary (Tyler and Heinbokel, 1985) and Gymnodinium
catenatum in Spanish Rias (Figueiras et al., 1995); blooms
of diatoms transitioning to dinoflagellates from wind-
induced upwelling of nutrient-rich deepwater in
mid-Chesapeake Bay (Malone et al., 1986; Sellner and
Brownlee, 1990); rainfall-induced fresher lenses of water
715
overlying salty, dense estuarine water (Loftus et al.,
1972); blooms of Microcystis in tidal fresh embayments
with little exchange (Linkov et al., 2007) or coastal
lagoons with limited flushing and long residence times
where raphidophyte or brown tide blooms are frequently
observed (Gobler et al., 2005; Handy et al., 2005); and
dense, water discoloring surface blooms of several dino-
flagellates and gas vesicle-rich cyanobacteria in highly
stratified and nutrient-poor surface waters (e.g., Sellner
and Brownlee, 1990; Suikkanen et al., 2007).
The latter bloom populations have remarkable diel
behaviors afforded to them through motility from
flagella and gas vesicle buoyancy, respectively. Diurnal
surface aggregation is assured through active upward
swimming of the dinoflagellates via positive phototaxis
(Kamykowski et al., 1998), while in the cyanobacterium,
flotation is assured through filling of thick-walled gas
vesicles in the cell, causing the cells to rise to the surface.
At night, the dinoflagellates may actively swim or pas-
sively settle to depth, while the cyanobacterium, now rich
in carbohydrate from daylong photosynthesis, sinks due
to the carbohydrate ballast that overwhelms the buoy-
ancy of the gas vesicles in the cyanobacterium. While at
depth, both groups assimilate nutrients. The dinoflagel-
lates respond to daylight again, actively swimming to
the surface. In contrast, nocturnal respiration of the car-
bohydrate in the cyanobacterium oxidizes the carbohy-
drate, removing ballast, and buoyancy produced
through gas-filled vesicles again returns the cyanobacte-
rium to the surface (e.g., Chu et al., 2007).
Finally, unique life cycles for some of the bloom-
forming species ensure continued growth opportunities
when light, nutrients, and physical conditions are
near optimal (see Fryxell, 1983). Many dinoflagellates
(e.g., Alexandrium, Gonyaulax, Protoceratium,
Peridinium, Diplopsalis, Diplopsalopsis, Diplopeltopsis,
and Scrippsiella) and raphidophytes (Heterosigma
akashiwo, Fibrocapsa japonica) produce temporary and
permanent resting stages (cysts) that on resuspension into
the overlying water column can excyst (burst from the
cyst wall) to resume the swimming behavior described
above. Similarly, some diatoms produce resting spores
(e.g., Chaetoceros, Leptocylindrus, Bacteriastrum),
enabling the same reemergence and growth possibilities.
For the cyanobacteria, some taxa (e.g., Aphanizomenon,
Anabaena) produce resting stages called akinetes and, like
cysts and spores, enable regrowth. Others (M. aeruginosa)
simply settle to the bottom in the fall to overwinter as near-
dormant populations on the sediment surface, only to be
resuspended in the late spring for reinitiating cell division
and accumulation into new blooms.
Summary
Toxic (and nontoxic) blooms of planktonic and benthic
single-cell organisms and accumulations of macroalgae
are increasingly common in the world’s nutrient-rich estuar-
ies, symptomatic of eutrophication attributable to mass land
716 TOXIC BLOOMS
use changes associated with human development. Toxic
blooms occur in tidal fresh and brackish portions of the estu-
ary, with a spectrum of toxicities on exposed estuarine fauna
as well as bird and mammal consumers: diarrhea, nausea,
and respiratory distress are common, while fish and shell-
fish mortalities can result from both exposure to toxin and
low dissolved oxygen concentrations associated with
high-biomass blooms of cyanobacteria, dinoflagellates,
raphidophytes, and macroalgae. Although many bloom-
forming species are favored in nutrient-rich estuaries, there
are several physical, physiological, and behavioral mecha-
nisms within estuaries and cells that can overlap to allow
growth and concentration of both toxic and nontoxic spe-
cies to ultimately adversely affect estuarine fauna and larger
consumers.
Bibliography
Anderson, D. A., Glibert, P. M., and Burkholder, J. M., 2002. Harm-
ful algal blooms and eutrophication: nutrient sources, composi-
tion, and consequences. Estuaries, 25, 704–726.
Chu, Z., Jin, X., Yang, B., and Zeng, Q., 2007. Buoyancy regulation
of Microcystis flos-aquae during phosphorus-limited and
nitrogen-limited growth. Journal of Plankton Research, 29,
739–745.
Figueiras, F. G., Wyatt, T., Alvarez-Salgado, X. A., and Jenkinson,
I. R., 1995. Advection, diffusion, and patch development in the
Rias Baixas. In Lassus, P., Arzul, G., Erard, E., Gentien, P.,
and Marcaillou, C. (eds.), Harmful Algal Blooms. Paris: Lavoi-
sier, Intercept, pp. 579–584.
Fistarol, G. O., Legrand, and Granéli, E., 2003. Allelopathic effect
of Prymnesium parvum on a natural plankton community.
Marine Ecology Progress Series, 255, 115–125.
Fistarol, G. O., Legrand, C., Selander, E., Hummert, C., Stolte, W.,
and Granéli, E., 2004. Allelopathy in Alexandrium spp.: effect
on a natural plankton community and on algal monocultures.
Aquatic Microbial Ecology, 35, 45–56.
Fryxell, G. A., 1983. Survival Strategies of the Algae. New York:
Cambridge University Press.
Gobler, C. J., Lonsdale, D. J., and Boyer, G. L., 2005. A review of
the causes, effects, and potential management of harmful brown
tide blooms caused by Aureococcus anophagefferens
(Hargreaves et Sieburth). Estuaries and Coasts, 28, 726–749.
Handy, S. M., Coyne, K. J., Portune, K. J., Demir, E., Doblin, M. A.,
Hare, C. E., Cary, S. C., and Hutchins, D. A., 2005. Evaluating
vertical migration behavior of harmful raphidophytes in the Del-
aware inland bays utilizing quantitative real-time PCR. Aquatic
Microbial Ecology, 40, 121–132.
Heisler, J., Glibert, P., Burkholder, J., Anderson, D., Cochlan, W.,
Dennison, W., Dortch, Q., Gobler, C., Heil, C., Humphries, E.,
Lewitus, A., Magnien, R., Marshall, H., Sellner, K., Stockwell,
D., Stoecker, D., and Suddleson, M., 2008. Eutrophication and
harmful algal blooms: a scientific consensus. Harmful Algae,
8, 3–13.
Kamykowski, D., Milligan, E. J., and Reed, R. E., 1998. Relation-
ships between geotaxis/photo-taxis and diel vertical migration
in autotrophic Dinoflagellates. Journal of Plankton Research,
20, 1781–1796.
Kemp, W. M., and Boynton, W. R., 1992. Benthic-pelagic interac-
tions: nutrient and oxygen dynamics. In Smith, D. E., Leffler,
M., and Mackiernan, G. (eds.), Oxygen Dynamics in the Chesa-
peake Bay A Synthesis of Recent Research. College Park, Mary-
land: Maryland Sea Grant Book, pp. 149–221.
Kim, C. S., Lee, S. G., Lee, C. K., Kim, H. G., and Jung, J., 1999.
Reactive oxygen species as causative agents in the
ichthyotoxicity of the red tide Dinoflagellate Cochlodinium
polykrikoides. Journal of Plankton Research, 21, 2105–2115.
Kubanek, J., Snell, T. W., and Pirkle, C., 2007. Chemical defense of
the red tide Dinoflagellate Karenia brevis against Rotifer graz-
ing. Limnology and Oceanography, 52, 1026–1035.
Linkov, I., Fristachi, A., Satterstrom, F. K., Shifrin, A., Steevens, J.,
Clyde, G. A., Jr., and Rice, G., 2007. Harmful cyanobacterial
blooms. In Linkov, I., Wenning, R. J., and Kiker, G. A. (eds.),
Managing Critical Infrastructure Risks, NATO Science for
Peace and Security Series C: Environmental Security. Nether-
lands: Springer, pp. 207–242.
Loftus, M. E., Subba-Rao, D. V., and Seliger, H. H., 1972. Growth
and dissipation of Phytoplankton in Chesapeake Bay. I. Response
to a large pulse of rainfall. Chesapeake Science, 13, 282–299.
Malone, T., Kemp, W. M., Ducklow, H., Boynton, W., Tuttle, J., and
Jonas, R., 1986. Lateral variation in the production and fate of
phytoplankton in a partially stratified estuary. Marine Ecology
Progress Series, 32, 149–160.
Marshall, J. A., de Salas, M., Oda, T., and Hallegraeff, G., 2005.
Superoxide production by marine microalgae. Marine Biology,
147, 533–540.
Nagle, D. G., and Paul, V. J., 1999. Production of secondary metab-
olites by filamentous tropical marine cyanobacteria: ecological
functions of the compounds. Journal of Phycology, 35,
1412–1421.
Pearson, L., Mihali, T., Moffitt, M., Kellmann, R., and Neilan, B.,
2010. On the chemistry, toxicology and genetics of the
cyanobacterial toxins, microcystin, nodularin, saxitoxin, and
cylindrospermopsin. Marine Drugs, 8, 1650–1680.
Prince, E. K., Myers, T. L., and Kubanek, J., 2008. Effects of harm-
ful algal blooms on competitors: allelopathic mechanisms of the
red tide Dinoflagellate “Karenia brevis”. Limnology and Ocean-
ography, 53, 531–541.
Sellner, K. G., and Brownlee, D. C., 1990. Dinoflagellate-
microzooplankton interactions in Chesapeake Bay. In Granéli,
E., Sundström, B., Edler, L., and Anderson, D. M. (eds.), Toxic
Marine Phytoplankton. New York: Elsevier, pp. 221–226.
Sellner, K. G., Doucette, G. J., and Kirkpatrick, G., 2003.
Harmful algal blooms: causes, impacts, and detection. Journal of
Industrial Microbiology and Biotechnology, 30, 383–406.
Suikkanen, S., Fistarol, G. O., and Granéli, E., 2004. Allelopathic
effects of the baltic cyanobacteria Nodularia spumdigena,
Aphanizomenon flos-aquae and Anabaena lemmermannii on
algal monocultures. Journal of Experimental Marine Biology
and Ecology, 308, 85–101.
Suikkanen, S., Laamanen, M., and Huttunen, M., 2007. Long-term
changes in summer Phytoplankton communities of the open North-
ern Baltic Sea. Estuarine Coastal Shelf Science, 71, 580–592.
Tang, Y. Z., and Gobler, C. J., 2010. Allelopathic effects of
Cochlodinium polykrikoides isolates and blooms from the Estu-
aries of Long Island, New York, on co-occurring Phytoplankton.
Marine Ecology Progress Series, 406, 19–31.
Tang, Y. Z., and Gobler, C. J., 2011. The green macroalga, Ulva
lactuca, inhibits the growth of seven common harmful algal
bloom species via allelopathy. Harmful Algae, 10, 480–488.
Tyler, M. A., and Heinbokel, J. F., 1985. Cycles of red water and
encystment of Gymnodinium pseudopalustre in the Chesapeake
Bay: effects of hydrography and grazing. In Anderson, D. M.,
White, A. W., and Baden, D. G. (eds.), Toxic Dinoflagellates.
New York: Elsevier, pp. 213–218.
Wang, D.-Z., 2008. Neurotoxins from marine Dinoflagellates:
a brief review. Marine Drugs, 6, 349–371.
Wang, R., Xiao, H., Zhang, P., Qu, L., Cai, H., and Tang, X., 2007.
Allelopathic effects of Ulva pertusa, Corallina pilulifera, and
Sargassum thunbergii on the growth of the Dinoflagellates
Heterosigma akashiwo and Alexandrium tamarense. Journal of
Applied Phycology, 19, 109–121.
 TRACE METALS IN ESTUARIES 717

Cross-references Since these comprise most of the elements in the periodic

Algal Blooms table, we should define those of concern in estuaries as
Anthropogenic Impacts those that are deposited from natural and anthropogenic
Cyanobacteria sources upriver. Typically, those of interest are of anthro-
Density Stratification pogenic origin, since they tend to be more evident down-
Dissolved Oxygen stream of industrial outfalls or mine drainages.
Estuarine Circulation
Eutrophication
Trace metals can be either essential for, or irrelevant to,
Flushing Time living organisms (i.e., nonessential). In regard to living
Macroalgae organisms, trace metals are typically defined as those that
Macroalgal Blooms are essential (Murray et al., 2009). However, in estuaries
Nutrients both essential and nonessential metals should be included
Oxygen Depletion in the definition since, in either case, they can be toxic in
Phytoplankton minute quantities – nanomolar concentrations or less,
Phytoplankton Blooms
Residence Time while those that are essential (micronutrients) are toxic
only when they substantially exceed metabolic needs –
typically, micromolar concentrations. These metals are
found in estuarine sediments, biota, and (to a lesser extent)
in the water column. Of these metals, two deserve special
TRACE METALS IN ESTUARIES attention. While most trace metals are found in living tis-
sues (i.e., they bioaccumulate), Hg and As* are capable
Peddrick Weis of biomagnification. This latter term means that they
Department of Radiology, Rutgers – New Jersey Medical increase, typically by an order of magnitude, with each
School, Newark, NJ, USA trophic level in the food web. This occurs because these
metals are turned into organometals (carbon-containing
Synonyms compounds) as a result of metabolic processes. Because
Heavy metals of the organic part of the molecule, an organometal can
passively cross the lipid bilayer that all cell membranes
Definition are made of, rather than requiring an active metabolic pro-
A metal is defined as an element that can form a salt by cess for uptake; they then are retained rather than excreted
replacing the hydrogen (H+) in an acid and form a base and are passed on to the next trophic level.
when combined with the hydroxyl (OH) radical. Organomercury compounds are particularly toxic. Some
organoarsenicals are toxic, while others are relatively
Description innocuous – e.g., the metabolism of inorganic As to
A few metals rank among the most common elements trimethylarsine renders it less toxic (Reimer et al., 2010).
on earth (Fe, Al, Ca, Mg), but most are less or much Table 1 summarizes the status of trace metals found in
less abundant. These can be considered trace metals. estuaries.

Trace Metals in Estuaries, Table 1 Trace metals in estuaries

Metal Symbol Essentialitya Capable of biomagnification Found in sediment Found in biota

Arsenicb As X X X
Beryllium Be X X
Cadmium Cd X X
Chromium Cr X X X
Cobalt Co X X X
Copper Cu X X X
Lead Pb X X
Manganese Mn X X X
Mercury Hg X X X
Molybdenum Mo X X X
Selenium Se X X X
Silver Ag X X
Vanadium V (X) X X
Zinc Zn X X X
a
While those metals listed as essential are as such for all eukaryotes and some bacteria, vanadium is known to be essential in relatively few
species, including tunicates
b
Arsenic is included here among “metals,” although it may be properly considered a “metalloid”
718 TROPHIC DYNAMICS
Bibliography
Murray, R. K., Bender, D. A., Botham, K. M., Kennelly, P. J.,
Rodwell, V. W., Weil, P. A., and Mayes, P. A., 2009.
Micronutrients: Vitamins & Minerals. Harper’s Illustrated
Biochemistry (28th ed.). New York: McGraw-Hill. Retrieved
January 4, 2012.
Reimer, K. J., Koch, I., and Cullen, W. R., 2010. Organoarsenicals.
Distribution and transformation in the environment. Metal Ions
in Life Sciences (Cambridge: RSC Publishing), 7, 165–229.
TROPHIC DYNAMICS
Xchel Moreno-Sánchez1, Andrés Abitia-Cárdenas1,
Juan M. Rodríguez-Baron2, Mónica Lara Uc2 and
Rafael Riosmena-Rodríguez2
1
Instituto Politécnico Nacional CICIMAR-IPN, La Paz,
BCS, Mexico
2
Programa de Investigación en Botãnica Marina,
Departamento Académico de Biología Marina,
Universidad Autónoma de Baja California Sur, La Paz,
Baja California Sur, Mexico
Synonyms
Trophic structure; Trophic web
Definition
Trophic dynamics is the basic process of transference
of energy from one trophic level to the next in an
ecosystem.
Introduction
The basic process of energy transfer in ecosystems occurs
through trophic dynamics (Lindeman, 1942). Trophic
dynamics in tropical and subtropical estuarine and coastal
marine ecosystems are characterized by a broad heteroge-
neity, which is determined by highly diverse biological
communities and complex trophic webs with a high
degree of interaction (Manickchand-Heileman et al.,
1998). The dynamic nature of trophic web interactions
and the variation in structure and function of several
coastal systems (including estuaries) are well known.
From the time of naturalists to the present-day ecolo-
gists, there has been an enormous interest in understand-
ing energy transfer in ecosystems, and research has
increased notably since Lindeman (1942) coined the con-
cept of trophic dynamics. This concept, along with the
articles published by Odum, fostered a rapid progress of
the use of the laws of thermodynamics in ecological
studies.
Estuarine trophic dynamics
Trophic dynamics have been widely studied in estuaries,
especially within the ichthyological community, but the
role of other biological components in the flow of energy
toward different trophic levels has only been suggested.
Estuaries are habitats with an exceptional primary produc-
tivity that allows them to support a high abundance and
biomass of fish; they are essential habitats in the life cycle
of several teleosts, functioning as feeding, reproduction,
and refuge areas. Fish communities in estuaries are very
dynamic due to seasonal changes in abiotic factors: cur-
rents, water temperature, pH, freshwater inputs, evapora-
tion, precipitation, and drastic changes in salinity. These
changing and extreme conditions result in estuaries being
characterized by few dominant fish species that are resi-
dent or typical of these areas. These species are classified
as anadromous (fish that migrate from salt water to fresh-
water to reproduce) or catadromous (fish that migrate from
freshwater to salt water to reproduce) (Kennish, 1990).
The trophic dynamics of estuaries are determined
mainly by the trophic relationships of all biological com-
ponents. The movement of nutrients between estuaries
and marine communities occurs through the biota in
a predictable sequence of trophic relations (nutrients).
The trophic structure of communities is centered on the
concept of the trophic web, which is the result of branched
and/or linked trophic chains. It can be defined as the sum
of all chains, with organisms grouped in categories or tro-
phic levels, which consist of producers, consumers, and
decomposers. These in turn are responsible for energy
transfer. Therefore, at the base of the trophic chain are all
the autotrophic organisms which fixate carbon through
photosynthesis and provide energy to the primary con-
sumers (heterotrophs) and so on through the upper trophic
levels. Primary consumers serve as food for secondary
consumers (herbivores), and these are a source of food
for tertiary consumers (carnivores). Decomposers
(saprophytic bacteria and fungi) assimilate dead plant
and animal matter, transforming it in organic matter to
get energy, while they release mineral nutrients that pri-
mary producers (autotrophs) use for growth. At each tro-
phic level, approximately 80–90 % of potential energy is
lost as heat, and this is a limiting factor that restricts tro-
phic chain lengths to three to four trophic levels.
Estuarine trophic dynamic processes
The feeding habits of fish are important for several rea-
sons. First, they reveal the trophic relations of the different
species and indirectly bring to light aspects of the energy
flow in estuarine communities. Second, they reveal the
ecological relations between organisms, which can be
used to better interpret the general dynamics of estuaries
and to make recommendations for the appropriate admin-
istration of fisheries resources. The functioning of estua-
rine and coastal marine ecosystems depends mostly on
using an external source of energy, solar radiation.
A portion of this incident energy is transformed by photo-
synthetic processes (primary production) carried out by
the phytoplankton (such as diatoms, dinoflagellates,
coccolithophorids, cyanophytes, and silicoflagellates)
 TROPHIC DYNAMICS
and phytobenthos (red, green, and brown algae)
(Lindeman, 1942). These components, along with detri-
tus, constitute the main energy source for organisms in
estuaries (Kennish, 1990). Zooplankton is comprised
mainly of crustaceans such as copepods, as well as mol-
luscs, coelenterates, and chaetognaths. The phytoplankton
is consumed by zooplankton; about 50 % of phytoplankton
is not consumed and goes into detritus. The zoobenthos
consists of a wide range of organisms, mostly invertebrates
such as sponges, crabs, echinoderms, polychaetes, snails,
clams, and sea stars, which feed on the phytobenthos as well
as on detritus. The zoobenthos is one of the main food
sources for fish, which compose the most conspicuous and
dominant community in estuaries. The abundance, biomass,
and diversity of the ichthyofauna are regulated by the size
of the estuary, the season (wet or dry), latitude (tropical
and subtropical zones), and changes in abiotic factors
(temperature, freshwater input, salinity changes, pH, currents
within the estuary, and evaporation). These factors can lead
to changes in the fish community structure through migration
or immigration of less eurytopic species. However, most of
the population is comprised by highly tolerant species that
have a clear tendency toward an r strategy (Pianka, 1970).
In general, four fish feeding categories can be observed
(Claro, 1994):
Herbivores: This trophic category includes fish that feed
exclusively on plants (algae). Fish from the Kyphosidae
family are very well represented in estuaries.
Detritivores: This group contains species that consume
detritus, which consists of vegetal and animal material
that has not been consumed and gets deposited on the
benthos. The most typical fish family in estuaries is
the Mugilidae family.
Benthophagous: Fish that feed on small organisms that
inhabit the substrate. Most fish species present in estu-
aries are found in this category (e.g., Mullidae and
Chaetodontidae families).
Carnivores: Consumers of animal prey, they feed mainly
on fish and invertebrates. The most representative spe-
cies belong to the families Lutjanidae, Serranidae,
Haemulidae, Carangidae, and Balistidae.
A practical way to analyze the trophic dynamics of estu-
aries is by performing functional diversity studies that pro-
vide a rapid characterization of communities, reducing
complex ecosystems into species groups (functional
groups) with ecological equivalence, thereby facilitating
comparative community studies (Root, 1967; Root, 2001).
Species that comprise a functional group are those that
overlap in the highest number of variables in the
multidimensional niche, being ecologically equivalent in
function (Nagelkerken and Van der Velde, 2004). From
the functional viewpoint, biological communities can be
represented as a mosaic of functional groups, or as bricks
with which communities are built (Krebs, 2003). The pres-
ence of multiple species (dominant and subordinates)
within each functional group increases functional
719
redundancy and contributes to the resilience of the estuary
or ecosystem studied (Hooper et al., 2005).
With a higher number of functionally similar species,
each with different responses to environmental factors,
there is a higher probability of at least one species surviv-
ing possible perturbations (Hooper et al., 2005). If there is
no functional redundancy, the loss of a single species
could result in the complete loss of a functional group;
therefore, at least one species per functional group is
essential for ecosystem functioning (Díaz et al., 2005).
Currently, due to urban and touristic development, anthro-
pogenic impacts affect negatively coastal areas and espe-
cially estuaries, so that immediate and continued
evaluations of these effects on the local fauna and flora
are extremely important. The study of functional diversity
is therefore an effective and precise tool to evaluate these
effects.
The most important inputs for the development and
evolution of trophodynamic perspectives over the last
decades have occurred mainly in the field of aquatic ecol-
ogy (Lindeman, 1942; Ulanowicz, 2004). The mass bal-
ance model ECOPATH with Ecosim has been the most
extensively used tool for analyzing trophic webs
(Polovina, 1984; Christensen and Pauly, 1992). This
model describes quantitatively the energy flows
(biomass) within food webs and facilitates an approxima-
tion to the structure and function of the ecosystem. Addi-
tionally, the model outputs allow the calculation of
several ecological indicators of the state of the trophic
web, such as ascendancy, surplus, and development poten-
tial (Ulanowicz, 1986; Libralato et al., 2006), that can be
used to follow up an ecosystem’s development through
time and to compare the ecological maturity of systems
located in different parts of the world.
Summary
Estuarine trophic dynamics is one of the most complex
processes of estuarine science. Its complexity lies in the
high environmental heterogeneity and biodiversity of
these coastal systems, which includes all trophic levels
from the smallest autotrophs to the largest predators. The
microbial decomposers must be considered as well.
Among the most important components in the flow of
energy in estuaries are fish communities due to the great
diversity of finfish eating habits which has resulted in
a large number of direct and indirect trophic interactions
with other biotic components present within networks that
structure and determine the energy dynamics of these
valuable ecosystems.
Bibliography
Claro, R., 1994. Características generales de la ictiofauna. In Claro, R.
(ed.), Ecologı́a de los peces marinos de Cuba. Capı́tulo 2. México:
Instituto de Oceanología y CIQRO, pp. 124–143.
Christensen, V., and Pauly, D., 1992. ECOPATH II: a software for
balancing steady-state models and calculating network charac-
teristics. Ecological Modeling, 61, 169–185.
720 TURBIDITY
Díaz, S., Tilman, D., Fargione, J., Chapin, F. S., III, Dirzo, R.,
Kitzberger, T., Gemmill, B., Zobel, M., Vilà, M., Mitchell, C.,
Wilby, A., Daily, G. C., Galetti, M., Laurance, W. F., Pretty, J.,
Naylor, R., Power, A., and Harvell, D., 2005. Biodiversity regu-
lation of ecosystem services. In Hassan, R., Scholes, R., and
Ash, N. (eds.), Ecosystems and Human Wellbeing: Current State
and Trends. Findings of the Condition and Trends Working
Group. Washington, DC: Island Press, pp. 297–329.
Hooper, D. U., Chapin, F. S., Ewel, J. J., Hector, A., Inchausti, P.,
and Lavorel, S., 2005. Effects of biodiversity on ecosystem func-
tioning: a consensus of current knowledge. Ecological Mono-
graphs, 75, 3–35.
Kennish, M. J., 1990. Ecology of Estuaries. Boca Raton: CRC
Press. Biological Aspects, Vol. II.
Krebs, C. J., 2003. Ecologı́a. Estudio de la Distribución y la
Abundancia, 2nd edn. México: Oxford University Press.
Libralato, S., Christensen, V., and Pauly, D., 2006. A method for
identifying keystone species in food web models. Ecological
Modelling, 195, 153–171.
Lindeman, R. L., 1942. The trophic-dynamic aspect of ecology.
Ecology, 23, 157–176.
Manickchand-Heileman, S., Soto, L. A., and Escobar, E., 1998.
A preliminary trophic model of the continental shelf, southwest-
ern Gulf of Mexico. Estuarine, Coastal and Shelf Science, 46,
885–899.
Nagelkerken, I., and van der Velde, G., 2004. A comparison of fish
communities of subtidal seagrass beds and sandy seabeds in
13 marine embayments of a Caribbean island, based on species,
families, size distribution and functional groups. Journal of Sea
Research, 52, 127–147.
Pianka, E., 1970. On the r- and K selection American. Naturalist,
105, 592–597.
Polovina, J. J., 1984. Model of a coral reef ecosystem. I: the
ECOPATH model and its application to French Frigate Schoals.
Coral Reefs, 3, 1–11.
Root, R. B., 1967. The niche exploitation pattern of the blue-gray
gnatcatcher. Ecological Monographs, 374, 317–350.
Root, R. B., 2001. Guilds. In Levin, S. A. (ed.), Encyclopedia of
Biodiversity. Durham, NC: Academic Press, pp. 295–302. 3.
Ulanowicz, R. E., 1986. Growth and Development: Ecosystem Phe-
nomenology. New York: Springer.
Ulanowicz, R. E., 2004. A synopsis of quantitative methods for eco-
logical network analysis. Computational Biology and Chemistry,
28, 321–339.
Cross-references
Food Web/Trophic Dynamics
TURBIDITY
Yuri Gorokhovich
Department of Earth, Environmental, and Geospatial
Sciences, Lehman College, City University of New York
(CUNY), Bronx, NY, USA
Turbidity
Turbidity is a measure of clarity in the water caused by
suspended matter, such as clay, silt, fine particulate
organic and inorganic matter, soluble colored organic
compounds, plankton, and other microscopic
microorganisms. Turbidity is an expression of the optical
property that causes light to be scattered and absorbed
rather than transmitted straight through a water sample.
Turbidity measurement is mandatory for regulatory
purposes.
In the middle of twentieth century, turbidity was mea-
sured using the Jackson candle turbidimeter. The method
was visual since the scientist evaluated light scattering
using unaided eyes. This method is no longer in use. The
modern method is called nephelometric, and it is based
on a comparison of the intensity of light scattered by the
sample under defined conditions with the intensity of light
scattered by a standard reference suspension (usually
formazin, a polymer) under the same conditions. Turbidity
units depend on the light source (e.g., white, near infra-
red), its angle in relation to the detector, and number of
detectors (i.e., ratio).
Turbidity is expressed in nephelometric turbidity units
(NTU) if the light source is white or broadband
(400–680 nm) located at 90 detection angle to one detec-
tor (Anderson, 2005), nephelometric turbidity ratio units
(NTRU) if the light source is at 90 detection angle and
measurement is done by multiple detectors with ratio com-
pensation, backscatter units (BU) if the light source is at
30  15 detection angle (backscatter), attenuation units
(AU) if the light source is at 180 detection angle
(attenuation), and nephelometric turbidity multibeam
units (NTMU) if there are multiple light sources and
detectors are located at 90 and possibly other angles to
each beam.
Sometimes turbidity can be expressed in formazin tur-
bidity units (FTU) comparable to NTU, obtained by
absorptiometric method measuring the amount of the
transmitted light absorbed by a formazin standard. This
method uses near-infrared (780–900 nm) or monochrome
light source. Depending on device, units can vary such as
FNRU (formazin nephelometric ratio units), FBU
(formazin backscatter units), FAU (formazin attenuation
units), and FNMU (formazin nephelometric multibeam
units) (Anderson, 2005).
Various studies made correlation between turbidity and
water quality parameters. The first study to use turbidity
for computation of suspended loads was presented by
Emmet (1975) and Truhlar (1976). Effler and Johnson
(1987) showed correlation between turbidity and calcium
carbonate precipitation in lakes. LeChevallier and Norton
(1992) showed that the removal of particles greater than
five microns and turbidity were useful predictors of
Giardia and Cryptosporidium removal. Strunk (1992)
identified correlation between turbidity and sewage
treatment plants and road discharge.
Bibliography
Anderson, C.W., 2005, Turbidity (ver. 2.1): U.S. Geological Survey
Techniques of Water-Resources Investigations, book 9, chap.
A6., sec. 6.7, September, 2005. Accessed May, 2013, from:
http://water.usgs.gov/owq/FieldManual/Chapter6/6.7_contents.
html. (May 27, 2013).
 TYCHOPLANKTON
Effler, S. W., and Johnson, D., 1987. Calcium carbonate precipita-
tion and turbidity measurements in Otisco Lake, New York.
Water Resources Bulletin, 23(1), 73–79.
Emmet, W. W., 1975. The Channels and Waters of the Upper
Salmon River Area, Idaho. USGS Professional Paper 870-A.,
115 p.
LeChevallier, M. W., and Norton, W. D., 1992. Examining relation-
ship between particle counts and Giardia, Cryptosporidium and
turbidity. Journal of the American Water Works Association,
84(12), 54–60.
Strunk, N., 1992. Case studies of variations in suspended matter
transport in small catchments. Hydrobiologia, 235(6), 247–255.
Truhla, J. F., 1976. Determining suspended sediment loads from tur-
bidity records. In Proceedings of the Third Federal Inter-Agency
Sedimentation Conference 1976, held at Denver, Colorado,
March 22–25, 1976. Water Resources Council, Sedimentation
Committee, pp. 7-65–7-74.
TYCHOPLANKTON
Lawrence Cahoon
Department of Biology & Marine Biology, University of
North Carolina Wilmington, Wilmington, NC, USA
Synonyms
“Tychopelagic” forms, in reference to diatoms (Hendey,
1964)
Definition
Hendey’s (1964) description of marine tychopelagic dia-
toms specified that these forms spend a major portion of
their life cycle attached to fixed substrata but become
planktonic after physical processes suspend them or tear
them loose from substrate. Hutchinson (1967), examining
freshwater forms, presented a broader definition of
tychoplankton, which included pseudoplankton (only
occasionally suspended in the water) and meroplankton
(spending a part of their life cycle in the plankton). Kuhn
et al. (1981) defined tychoplanktonic species as occupying
benthic niches but under certain conditions proliferating in
the plankton. The utility of distinguishing tychoplankton
includes studies of past conditions (paleoecology).
721
Taxonomy
Most published references to tychoplankton refer to dia-
toms that are typically found on the bottom or other solid
substrate and that enter the water column when physical
forces (waves, tidal currents) suspend them. Owing to
the difficulty of distinguishing tychoplankton from obli-
gately benthic or planktonic forms except by identification
to species and counting, estimates of tychoplankton bio-
mass and production are challenging, although
tychoplankton are likely ubiquitous in estuarine ecosys-
tems. Inclusion of meroplankton, species regularly spend-
ing part of their life cycle in the plankton, with the
tychoplankton is not a common usage in the literature on
estuarine ecology.
Example species
Vos and DeWolf (1993) identified the diatoms Cymatosira
belgica and Raphoneis minutissima as tychoplankton spe-
cies. Espinosa et al. (2006) identified the diatoms Paralia
sulcata, Staurosira construens, and Staurosirella pinnata
as tychoplankton.
Bibliography
Espinosa, M. A., Hassan, G. S., and Isla, F. I., 2006. Diatom distri-
bution across a temperate microtidal marsh, Mar Chiquita coastal
lagoon, Argentina. Thalassas, 22, 9–16.
Hendey, N. I., 1964. An Introductory Account of the Smaller Algae
of British Coastal Waters. Part V. Bacillariophyceae (Diatoms).
Ministry of Agriculture, Fisheries and Food, Fishery Investiga-
tion Series IV.
Hutchinson, G. E., 1967. A Treatise on Limnology. Volume
II. Introduction to Lake Biology and Limnoplankton. New York:
Wiley.
Kuhn, D. L., Plafkin, J. L., Cairns, J., Jr., and Lowe, R. L., 1981.
Qualitative characterization of aquatic environments using dia-
tom life-form strategies. Transactions of the American Micro-
scopical Society, 100, 165–182.
Vos, P. C., and de Wolf, H., 1993. Diatoms as a tool for
reconstructing sedimentary environments in coastal Wetlands.
Hydrobiologia, 269(270), 285–296.
Cross-references
Phytoplankton
Wave-Driven Sediment Resuspension