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Fluid dynamics for ecologists

Fluid dynamics play a crucial role in the study of marine ecosystems, and their understanding is essential for ecologists conducting scientific research in these environments. Waves and currents impact coastal habitats directly, influencing their physical structure, flow patterns, shear bottom stress and substrate stability, crucial for species distribution. Additionally, hydrodynamics govern transport processes like sedimentation, nutrient cycling, and larval dispersal, fundamental for ecosystem function. They also shape biological interactions, affecting behaviors, feeding strategies, and community dynamics of marine organisms. Understanding wave dynamics is crucial for assessing coastal resilience, predicting critical thresholds, and identifying conservation priorities in the face of climate change and human disturbances.

Fluid dynamics is the study of the movement of fluid. Among other things, it addresses velocity, acceleration, and forces exerted by or upon fluids in motion. In this blog post, a review is provided of the basic concepts of fluid dynamics that affect the life of macrophytes. Linear wave theory, wave propagation and benthic boundary layers are summarised.

Waves

Water flow in coastal marine ecosystems can be divided into unidirectional and oscillatory flows. Unidirectional flows (also described as currents) are those in which water particles tend to move in the same direction over time. These flows in marine ecosystems are mainly generated by tides. Oscillatory flows (also described as waves) are those in which water particles move in two directions at a periodic interval. These flows are generated by waves. Depending on the dominance of wave periodicity, a marine ecosystem can be characterized as tide-dominated or wave-dominated. This blog post is mainly focused on wave-dominated flows, and for simplicity, the currents and tides characteristics will not be reviewed in this post.

Wind generated waves are formed by the frictional stress produced by two fluid layers of different speed, which creates a transfer of energy from the wind to the water surface. Wind-generated waves are surface waves that occur on the ocean, sea, lake, rivers, and canals or even on small puddles and ponds. Wave size depends on the wind speed blowing over a distance of sea surface (known as wind fetch) for a lenght of time. Here is a useful online wave calculator to estimate wave conditions using known wind speeds and fetch, but remember that bathymetry and bottom slope will affect the propagation of waves too as you will see below.

Wave data can be obtained by direct measurements of the sea conditions using oceanographic instruments or estimated using numerical models. Check an early post on instrument deployments at sea. Oceanographic instruments are used to measure waves such as pressure sensors, accelerometers, acoustic dopplers, radars or altimeters mounted on satellites. Instruments can be deployed anchored at the sea bottom or fixed at meteorological buoys or at the shore. Wave records are generally very limited in space and time due to the cost of the instruments and measuring platforms. Also, waves are usually recorded on a specific location over a period of time. For these reasons, numerical models are used to estimate the wave conditions in locations where wave measurements are not available. In order to obtain a wave field over larger portions of the sea surface area where wave measurements are not available, numerical models are used. Numerical models are widely used, for example, to estimate wave fields over the Mediterranean Sea (Cañellas et al. 2007), to study a specific coastal area with an ecological, economical or turistic interest (Álvarez-Ellacuria et al. 2009, Álvarez-Ellacuria et al. 2010).

Waves are characterized by wave height H (from trough to crest) or wave amplitude a (half of the wave height), wavelength L (from crest to crest), wave period T (time interval between arrival of consecutive crests at a stationary point) and wave propagation direction (Fig. 1). Waves in a given area typically have a range of heights. For weather reporting and for scientific analysis of wind wave statistics, their characteristic height over a period of time is usually expressed significant wave height, Hs, which represents an average height of the highest one-third of the waves in a given time period. The angular frequency is ω = 2π/T, and the wave number is k = 2π/L.

Airy wave theory or linear wave theory gives a linear system description of the wave propagation of gravity waves on the surface of a homogeneous fluid layer. The theory assumes that the fluid layer has a uniform mean depth and that the fluid flow is inviscid, incompressible, and irrotational. Linear wave theory uses a potential flow approach to describe the motion of gravity waves on a fluid surface. In nature, a combination of waves and currents is present, but for simplicity, we deal with them independently.

Wave characteristics, shoaling, wave length, wave height, depth

Figure 1. Diagram of a shoaling wave approaching to shore. Wave height (H), wave length (L), water depth (d) and wave celerity or speed (C).

Assuming linear wave theory, the dispersion relationship in deep and shallow water describes the field of propagating waves as
\[ \omega^2 = gk \tanh(kh) \]  [1]
where g is the acceleration of gravity and h is the water depth. Substituting ω and K in Eq. 1 can also be writen as
\[ L = \frac{T^2 g}{2\pi} \tanh\left(\frac{2\pi h}{L}\right) \]  [2]
Waves approaching the coast travel through different water depths and are classified into three regimes; deep, shallow and deep waves. The speed of wave propagation, C,  can be expressed as
\[ C^2 = \frac{g}{k} \tanh(kh) \]  [3]
In deep water, kh is large and tanh(kh) = 1, therefore L = T^2g/2π or C = pg/k (Fig. 2). For a water depth larger than half the wavelength, h > L/2, the orbits are circular, and the phase speed of the waves is hardly influenced by depth (this is the case for most wind waves on the sea and ocean surface). In shallow water, kh is small and tanh(kh) = kh, therefore
\[ C = \sqrt{gh} \] [4]
For a water depth smaller than the wavelength divided by 20 (h < L/20), the orbits become elliptical or flatter due to the influence of the bottom. The phase speed of the waves is only dependent on water depth, and is no longer a function of periodic function or wavelength. All other cases, L/20 < h < L/2, are termed intermediate waves, where both water depth and period (or wavelength) have a significant influence on the solution of Airy wave theory. In this case, waves combine characteristics of deep and shalow water waves and speed is described as Eq. 3).
Shallow waves, intermediate waves, deep waves

Figure 2. Transition between shallow, intermediate and deep waves

Assuming linear wave theory, deep and shallow waves differ in characteristics other than wave velocity. In deep-water waves, the individual particles describe circles whose radius decreases with depth. At the surface, the radius is the same as the wave amplitude, and the particle velocity is the circumference of the circle divided by the wave period. In shallow-water waves, the particles describe ellipses. The radius along the minor axis is equal to the wave amplitude at the surface and decreases linearly with depth. At the bottom, the minor axis is zero, and the motion is horizontal. The radius of the major axis is a function of water depth, wavelength and amplitude. Here, near-bottom orbital velocity, ub, is given by
\[ u_b = \frac{2 \pi a b}{T} \]   [4]

where ab is the wave orbital amplitude calculated as

\[ ab = \frac{H_s}{2 \sinh\left(\frac{2\pi h}{L}\right)} \] [5]

where L is calculated iteratively from Eq 2.

Orbital velocities can also be measured from instruments deployed in the seafloor. Records from acoustic doppler velocimeters (ADV) provide large datasets to calculate velocity and turbulence. Field records are usually filtered to remove spikes and low beam correlations. Then velocity data can be filtered with a low pass filter to remove high frequency Doppler noise (Lane et al. 1998). Horizontal flow velocity is calculated as the root mean square (rms).

\[ U_{\text{rms}} = \sqrt{\frac{1}{N} \sum_{i=1}^{N} x_{i}^{2} + y_{i}^{2}} \] [6]

where x and y are the two horizontal velocities.

Wave energy possessed by a wave is in two forms, kinetic and potential. Kinetic energy is the energy inherent in the orbital motion of the water particles, while potential energy is possessed by the particles when they are displaced from their mean position. The total wave energy E per unit area is

\[ E = \frac{1}{8} \rho g H_{s}^{2} \] [7]

Wave propagation

As water waves propagate from the region where they were generated to the coast, both wave amplitude and wavelength are modified due to refraction (e.g., changes in bathymetry or interactions with wind-induced currents), diffraction (e.g., intense variations of the bottom), the loss of energy due to shoaling, wave damping and finally wave breaking. The surf zone is a highly dynamic area where energy from waves is partially dissipated through turbulence in the boundary layer and transformed in short and long waves, mean sea level variations and currents. Therefore, the energy dissipated in the surf zone is used for sediment transport providing a highly variable morphological environment (Dean and Dalrymple 2002).

Early efforts to study wave transformation from deep to shallow water were based on the geometrical ray theory. As a modification to linear wave theory, the mild slope approach, which assumes that the water depth changes slowly in a typical wave length, appears as an improvement over the former since it includes wave diffraction. In this approach, the vertical structure of the velocity is the same as that for a progressive wave over a constant depth, with the governing partial differential equations of the elliptic type. Moreover, these equations can be easily converted into parabolic type by assuming that the wave amplitude is primarily a function of water depth due to shoaling. This approach, known as the parabolic approach, can be seen as a modification of the ray theory where wave energy can be diffused along the rays as wave propagate (Liu and Losada 2002).

Boundary layers

The purpose of studying benthic boundary layers in ecology is to understand the water movement around organisms that live on or near the substratum. The boundary layer is the layer of fluid in the immediate vicinity of a bounding surface where effects of fluid viscosity are considered in detail. The boundary layer effect occurs in the field region where all changes occur in the flow pattern. The boundary layer distorts surrounding non-viscous flow. It is a phenomenon of viscous forces. This effect is related to the Reynolds number. The thickness of the boundary layer, δ, is normally defined as the distance from the solid body at which the flow velocity is 99% of the free stream velocity, which is the velocity calculated at the surface of the body in an inviscid flow solution.

When benthic marine macrophytes are present, the boundary layer is modified by the canopy, which influences the mean velocity, turbulence and mass transport (Nepf and Vivoni 2000, Ghisalberty and Nepf 2002, Luhar et al. 2010). Macrophytes reduce the flow velocity near the bottom, altering the logarithmic velocity profile (Fig. 3).

Vertical velocity of flow profile and vegetation

Figure 3.  Vertical profile of flow velocity interacting with a seagrass meadow. Top, weak currents (black line) and fast currents (dashed line). Bottom, velocity reduction due to benthic macrophytes.

Water flow can either be smooth and regular as the fluid flows in layers (laminar flow) or rough and irregular (turbulent flow). This depends on the velcity and the length scale (temporal and spatial scales, respectively) and is defined by the Reynolds number. The Reynolds number Re is a dimensionless number that gives a measure of the ratio of inertial forces ρν2/l∗ to viscous forces μν/l∗2 and consequently quantifies the relative importance of these two types of forces for given flow conditions, and is expressed as
\[ \text{Re} = \frac{u \cdot l_*}{\nu} \] [8]
where u is the flow velocity, l∗ is the characteristic length, and ν is the kinematic water viscosity.
Macrophytes are a source of drag; they reduce near-bottom water flow and dissipate current and wave energy. Drag forces act in a direction opposite to the oncoming flow velocity. Drag on macrophytes increases with water velocity and foliar surface area. In flexible organisms, foliar surface area will change with increasing flow velocity, and this change is encapsulated in the drag coefficient. The drag force is expressed as
\[ F_D = \frac{1}{2} \rho A f C_D u^2 \] [9]
 
where ρ is density of the water, Af is the frontal area, CD is the drag coefficient and u is the water velocity. The drag coefficient CD is a dimensionless parameter used to quantify the drag or resistance of an object in a fluid environment. It is used in the drag equation, where a lower drag coefficient indicates the object will have less hydrodynamic drag. The drag coefficient is always associated with a particular surface area.

Related research articles

Acoustic Doppler Velocimeter (ADV) Vectrino, Nortek measuring wave action and flow velocities in a Posidonia seagrass meadow.

6. Effect of a seagrass (Posidonia oceanica) meadow on wave propagation

Journal Papers
Infantes E, Orfila A, Simarro G, Luhar M, Terrados J, Nepf H
Marine Ecology Progress Series 456: 63-72
Publication year: 2012
Edge of Posidonia oceanica seagrass meadow on sandy bottom in Cala Millor, Mallorca Island, Spain, Mediterranean Sea.

1. Wave energy and the upper depth limit of Posidonia oceanica

Journal Papers
Infantes E, Terrados J, Orfila A, Cañellas B, Álvarez-Ellacuria A
Botanica marina 52: 419-427
Publication year: 2009
Artificial seagrass on hydraulic flume at MIT

3. Wave induced velocities inside a model seagrass bed

Journal Papers
Luhar M, Coutu S, Infantes E, Fox S, Nepf H
Journal of Geophysical Research Vol. 115, C12, 1-15
Publication year: 2010

14. Seagrass blade motion under waves and its impact on wave decay

Journal Papers
Luhar M, Infantes E, Nepf H
Journal of Geophysical Research - Oceans, 122.
Publication year: 2017
Effect of eelgrass (Zostera marina) on wave attenuation at Lomma Bay

Effect of eelgrass (Zostera marina) on wave attenuation at Lomma Bay

Reports
Infantes E
County Administrative Board of Skåne (Länsstyrelsen Skåne) Report 2020:07. ISBN: 978-91-7675-186-2
Publication year: 2020

34. Making realistic wave climates in low-cost wave mesocosms: a new tool for experimental ecology & biogeomorphology

Journal Papers
Infantes E, de Smit J, Tamarit E, Bouma TJ
Limnology and Oceanography: Methods, 19: 317-330
Publication year: 2021

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