Scholarly article on topic 'Assessing the impacts of sediments from dredging on corals'

Assessing the impacts of sediments from dredging on corals Academic research paper on "Earth and related environmental sciences"

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Marine Pollution Bulletin
{Corals / Dredging / Turbidity / Light / Thresholds / "Water quality"}

Abstract of research paper on Earth and related environmental sciences, author of scientific article — Ross Jones, Pia Bessell-Browne, Rebecca Fisher, Wojciech Klonowski, Matthew Slivkoff

Abstract There is a need to develop water quality thresholds for dredging near coral reefs that can relate physical pressures to biological responses and define exposure conditions above which effects could occur. Water quality characteristics during dredging have, however, not been well described. Using information from several major dredging projects, we describe sediment particle sizes in the water column/seabed, suspended sediment concentrations at different temporal scales during natural and dredging-related turbidity events, and changes in light quantity/quality underneath plumes. These conditions differ considerably from those used in past laboratory studies of the effects of sediments on corals. The review also discusses other problems associated with using information from past studies for developing thresholds such as the existence of multiple different and inter-connected cause-effect pathways (which can confuse/confound interpretations), the use of sediment proxies, and the reliance on information from sediment traps to justify exposure regimes in sedimentation experiments.

Academic research paper on topic "Assessing the impacts of sediments from dredging on corals"


MPB-07265; No of Pages 21

Marine Pollution Bulletin xxx (2015) xxx-xxx

Contents lists available at ScienceDirect Marine Pollution Bulletin journal homepage: Si MABIWE POIXOTTBON BumxETiM


Assessing the impacts of sediments from dredging on corals

Ross Jones a,e'*, Pia Bessell-Browne a,b,e, Rebecca Fisher a,e, Wojciech Klonowskic,d, Matthew Slivkoffc,d

a Australian Institute of Marine Science (AIMS), Perth, Australia

b Centre of Microscopy, Charaterisation and Analysis, The University of Western Australia, Perth, Australia c Curtin University, Bentley, Perth, Australia d In situ Marine Optics, Bibra Lake, Perth, Australia e Western Australian Marine Science Institution, Perth, Australia


There is a need to develop water quality thresholds for dredging near coral reefs that can relate physical pressures to biological responses and define exposure conditions above which effects could occur. Water quality characteristics during dredging have, however, not been well described. Using information from several major dredging projects, we describe sediment particle sizes in the water column/seabed, suspended sediment concentrations at different temporal scales during natural and dredging-related turbidity events, and changes in light quantity/quality underneath plumes. These conditions differ considerably from those used in past laboratory studies of the effects of sediments on corals. The review also discusses other problems associated with using information from past studies for developing thresholds such as the existence of multiple different and inter-connected cause-effect pathways (which can confuse/confound interpretations), the use of sediment proxies, and the reliance on information from sediment traps to justify exposure regimes in sedimentation experiments. Crown Copyright © 2015 Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license



1. Introduction..............................................................................................................................0

2. Natural and dredging-related turbidity events................................................................................................0

3. Conceptual model of the effects of sediments on corals........................................................................................0

4. Cause-effect pathways....................................................................................................................0

4.1. Suspended-sediment................................................................................................................0

4.2. Light availability....................................................................................................................0

4.3. Sediment covering..................................................................................................................0

4.4. Combined effects of proximal stressors................................................................................................0

5. Water quality characteristics of dredging plumes............................................................................................0

5.1. Particle size distributions (PSDs) associated with dredging..............................................................................0

5.2. Suspended-sediment concentrations (SSCs)..........................................................................................0

5.3. Light attenuation and light quality....................................................................................................0

6. Experimental studies of the effect of sediments on corals......................................................................................0

6.1. Suspended-sediments..............................................................................................................0

6.2. Sediment deposition................................................................................................................0

7. Discussion................................................................................................................................0

7.1. Experimental studies of the effects of suspended-sediment and light attenuation on corals................................................0

7.2. Experimental studies of the effects of sediment deposition on corals....................................................................0

7.3. Seawater quality thresholds for coral reefs and future directions........................................................................0

Author contributions............................................................................................................................0

Competing interests............................................................................................................................0



* Corresponding author at: Australian Institute of Marine Science (AIMS), Perth, Australia. E-mail address: (R. Jones).

0025-326X/Crown Copyright © 2015 Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (

Article history:

Received 16 September 2015 Received in revised form 19 October 2015 Accepted 22 October 2015 Available online xxxx





Thresholds Water quality


2 R. Jones et al. / Marine Pollution Bulletin xxx (2015) xxx-xxx

1. Introduction

Dredging and dredging related activities such as dredge material placement (spoil disposal) releases sediment into the water column where it can have a significant effect on nearby benthic communities (Bak, 1978; Brown et al., 1990; Dodge and Vaisnys, 1977; Erftemeijer et al., 2012; Foster et al., 2010; Rogers, 1990). Consequently, most large-scale dredging projects require environmental impact assessments (EIAs) and active management when underway (EPA, 2011). The environmental and management issues associated with dredging and construction around ports and harbours have recently been reviewed and guidelines produced for monitoring, management, and mitigation (Foster et al., 2010).

There is a critical need to improve the ability to make scientifically sound predictions of the likely extent, severity, and persistence of environmental impacts associated with dredging especially when conducted close to sensitive habitats such as coral reefs. One of the more practical ways is by water quality monitoring, i.e. measuring the key hazards which are capable of having adverse biological effects. Water quality monitoring can be vessel-based or by fixed or rapidly relocatable mobile 'sentinel' in situ platforms, or even autonomously by vehicles remotely controlled from land (Van Lancker and Baeye, 2015). The use of these technologies allows dredging proponents to be quickly alerted to conditions where environmental damage may occur and modify the dredging activities as required. However, to be useful in this way, the water quality data have to be coupled to thresholds that relate the physical pressure(s) to a biological response(s) and define exposure conditions above which effects could occur.

Similar threshold values for chemical contaminants are usually derived from field data and more frequently from controlled, laboratory-based studies (i.e. exposure studies) where test organisms are incubated with toxicant(s)/hazards over short-term (acute) and longer term (chronic) time periods (Chapman, 2002). Many such studies have been conducted on the effects of sediments on corals in the context of understanding the effects of terrestrial run-off and dredging, and the studies have been collated and reviewed many times(i.e., Erftemeijer et al., 2012; Fabricius, 2005; Foster et al., 2010; Jones et al., 2015b; Rogers, 1990). These studies have contributed to our collective understanding and concern of the environmental effect of sediments on coral reefs. Logically, information from these studies should be suitable for threshold development. However, the results are dependent on the exposure scenarios and experimental conditions used, and it is noticeable that although a great deal of data have been collected (see, Falkenberg and Styan, 2014), there is little information in the public literature about the conditions that can occur in situ during dredging around coral reefs (Jones et al., 2015a). Recently, Harris et al. (2014) highlighted the fundamental significance in ecotoxicology of the use or environmentally relevant or realistic conditions, the need for comprehensive justification of any exposure conditions, and the dangers associated with an incomplete understanding.

We first identify, through a conceptual model, the likely exposure pathway(s) associated with the effects of dredging, including how proximal stressors are interlinked and interact with natural stressors. Data from several recent major dredging programmes in reefal areas of tropical Australia are then examined to describe how the water quality conditions change over time to provide a first order approximation of the environmentally relevant or realistic exposure scenarios. The existing literature of the effects of sediments on corals is then examined with specific attention to the experimental conditions used in order to assess the suitability of past studies for deriving information for impact prediction purposes (i.e. for modelling and monitoring). One of the objectives of the review is also to facilitate a technical understanding of sediments and sediment resuspension, water quality issues associated with dredging programmes, and some of the terminologies used and processes involved. The analysis concludes with some recommendations and guidance for future study.

2. Natural and dredging-related turbidity events

Turbidity is the optical property of a suspension that causes light to be scattered and absorbed rather than transmitted through the water column (Davies-Colley and Smith, 2001); it implies muddiness and lack of clarity and transparency (Kirk, 1994). The attenuation of light is related to the suspended-sediment concentration and the water itself, plankton, and other microscopic organisms and coloured organic matter (humics, Kirk, 1994). Suspended-sediments are often the dominant factor affecting light attenuation in tropical and sub-tropical waters.

Natural turbidity events are common in the shallow reef environment and although resuspension and transport of suspended material may be strongly influenced by unidirectional currents, wind-driven waves are the primary mechanism of turbidity generation (Jing and Ridd, 1996; Larcombe et al., 2001; Lawrence et al., 2004; Ogston et al., 2004; Verspecht and Pattiaratchi, 2010). The underlying hydrodynamic principles associated with natural turbidity generation are illustrated in Fig. 1 and further information on hydrodynamic controls of sediment resuspension and sedimentation are given in Larcombe and Woolfe (1999) and in a suite of more recent studies in Hawaii (Ogston and Field, 2010; Ogston et al., 2004; Storlazzi and Jaffe, 2008; Storlazzi et al., 2004).

Briefly, water moves in circular motion with the diameter oforbit equal to the wave height (Van Rijn et al., 1990) and the orbital motion is transferred into deeper water with the radius of the motion decreasing with increasing depth and compressing into ellipses in shallow water (Perry and Taylor, 2009). When tidally-driven near-bottom currents and wave-orbital nearbed speeds are sufficient to exceed a critical bed shear stress seabed erosion occurs, releasing sediment into the overlying column as a suspended load (Ogston et al., 2004; Storlazzi et al., 2004; van Rijn, 2007). Once sediment is suspended a small rise in shear stress can result in a disproportionate increases in suspended-sediment concentration (SSCs) (Orpin et al., 2004). Sediments will remain in suspension until the fluid velocity is insufficient for turbulent eddies to balance gravitational forces and the particles will settle out, depositing on the seabed (Masselink et al., 2014). This could occur in quiescent, calm areas (i.e. embay-ments), in waning periods after storms (Storlazzi et al., 2009; Verspecht and Pattiaratchi, 2010) or during slack water between tides. Fluctuating shear stresses can also result in successive resuspension and deposition events and if the amount of sediment deposited on the bed exceeds the amount eroded over the same time period then net deposition will occur (McAnally and Mehta, 2001; Ogston et al., 2004).

The phrase 'turbidity-generating event' is used here generically to describe dredging and marine construction related activities which release sediments into the water column. These include release of sediment at the seabed by the action of the drag head or cutter suction head (Fig. 2), release of sediments at the surface by the overflow of fine material from the dredge hopper or hopper barge, spillage from the bucket of a back hoe dredge, and also dredge material disposal. Release from the overflow is one of the primary mechanisms and sediment concentrations in the hopper can measure in the tens of g L-1 range; however, there is usually an abrupt, rapid initial dilution by a factor of 10-100, with SSCs decreasing logarithmically with time and distance (Duclos et al., 2013; Spearman, 2015; Spearman et al., 2007).

The terms 'near-field' and 'far-field' are used to distinguish sediment plumes based on the behaviour of the resuspended particles and this is closely related to the proximity to dredging (for a detailed discussion see VBKO (2003)). Near-field plumes are also called 'active' plumes and behave in a 'dynamic' manner, with material descending rapidly to the seabed as a density current. Entrainment of ambient water into the plume can slow its descent and entrainment of air bubbles into the plume on discharge and action from the ship's propellers can also lift a portion of the plume to the surface (Fig. 2, see also Jones, 2011b).


Fig. 1. Stylized representation of current and wave-induced turbidity generation in the shallow, reef environment (see text for explanation).

Settling of coarser rather than finer particles occurs as sediment is advected away from the dredging site and sands will drop quickly within the first 10-20 min (Duclos et al., 2013). The far-field or passive plume is made up of much lower SSCs and the fine fraction of material mixed into the water column can stretch for many kilometres and persist for several hours, as determined by site-specific hydrodynamics and local conditions (Fig. 3B).

3. Conceptual model of the effects of sediments on corals

A conceptual model of the effects of turbidity-generating events on corals based on the US Environmental Protection Agency (USEPA) causal/diagnosis decision information system framework (CADDIS)(Norton et al., 2009; USEPA, 2004) is shown in Fig. 4. In the model, all known cause-effect linkages, biologically plausible cause-effect pathways, and potentially confounding variables and interacting factors are displayed in a single framework. Sources of sediments include carbonate and igneous and siliciclastic intertidal and subtidal sediments released into the water column (Fig. 4). There are many activities associated with dredging and land reclamation activities that can cause turbidity ranging from major sources such as hopper overflow to more trivial sources such as rockdumping (Koskela et al., 2002). Damage associated with dredging and turbidity-generating activities can be separated into direct and indirect effects (Fig. 4), with direct effects including the removal of hard and soft substrate (within the dredge footprint), and smothering of the seabed at disposal (placement) sites.

A second and much larger group of cause-effect pathways are associated with mobilization of sediments into the water column, i.e. 'turbidity' or 'plume-generation' and subsequent movement of sediment out of the immediate dredging or dredge material placement site.

These 'indirect' effects can be further divided into (1) chemical and (2) physical effects. Chemical effects include the release of FeS-rich sediments which can rapidly deoxygenate water and change pH levels (i.e. a proximal stressor) (Morgan etal., 2012). Desorption of legacy contaminants and release of nutrients and pore-water dilution can also change the chemical environment of the water column (Eggleton and Thomas, 2004; Su et al., 2002). Nutrient release has the potential to change oxygen concentrations, mediated through phytoplankton and microbial blooms (i.e. a step in the causal pathway, Fig. 4). Changes in oxygen and nutrient levels and especially contaminant concentrations have the potential for acute and chronic toxicological, cellular and physiological effects, including genotoxic (mutagenic, teratogenic and carcinogenic) effects, as well as bioaccumulative effects through uptake and ingestion of contaminants (see for example Hedge et al. (2009)). Prior to dredging, sediments are normally examined for contaminant concentrations and, in Australia at least, landfilled if levels exceed screening guidelines (DEWHA, 2009). Many capital dredging projects in the tropics also occur in green-field sites without historical pollution, and sediment contamination is typically less of a concern than industrialized areas (with a few notable exceptions, see Jones, 2011a). Whilst not discounting the potential significance of chemical effects, especially in the near-field environment, the rest of the review is concerned with indirect effects of physical proximate stressors associated with turbidity generating events and include (1) elevated suspended-sediments, (2) changes in light quality and quantity, and (3) sediment covering (Fig. 4).

For all proximate stressors the key interacting factors are periods of naturally elevated turbidity associated with (1) currents, wind-driven waves and elevated sea states associated with trade winds and storms (Jing and Ridd, 1996; Larcombe et al., 2001; Lawrence et al., 2004; Ogston et al., 2004; Orpin and Ridd, 2012; Verspecht and Pattiaratchi,

Fig. 2. Stylized representation of plume generation by a trailing suction hopper dredge, the most commonly used of the hydraulic dredges for soft sediments (see Foster et al., 2010). Turbidity is generated at the drag-head and at the surface if sediment-laden water inside the hopper is allowed to discharge (overflow). This can occur from the ship's sides but usually through a vertical shaft inside the hopper, exiting below water level (Foster et al., 2010; VBKO, 2003).


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Fig. 3. (A) Working trailing suction hopper dredge in the Pilbara region ofWestern Australia generating buoyant surface plumes through propeller wash. (B, C) Atmospherically corrected, colour corrected, pan sharpened satellite image from the United States Geological Survey (USGS) Operational Land Imager (OLI) instrument on (B) 10 July 2014, showing sediment plumes caused by dredging and dredge material placement near Onslow in the Pilbara region ofWestern Australia (where DMPS = offshore dredge material placement site) and (C) 25 September 2013, showing natural turbidity caused by a high wind event. (D) Buoyant turbidity plume overlying a branching Acropora spp. colony. (E) Abranching Acropora spp. colony photographed during the daytime, and temporarily in darkness caused by the elevated turbidity levels. (F) Smothering of a plating Montipora spp. by sediment. Images D and F were taken during a dredging project at Magnetic Island (Central Great Barrier Reef) and image E during the Barrow Island project. Satellite images courtesy of Mark Broomhall and Peter Fearns (Department of Imaging and Applied Physics Curtin University of Technology).

2010) and (2) in the nearshore, coastal environment, the effects of river-borne discharges from monsoonal events and tropical depressions, and associated sediment-laden, hypopycnal flood waters (Orpin and Ridd, 2012; Storlazzi et al., 2009). Natural cycles of light availability are also an interacting factor and summarized in Anthony et al. (2004)), these include (1) the seasonal pattern of daily surface irradiance (insolation) governed by the solar declination cycle (Kirk, 1994), (2) tidal cycles which affect the depth of the water, (3) meteorological phenomena including patterns of cloud formation (Wright, 1997) and (4) large-scale pressure systems such as the Madden-Julian oscillation (MJO) (Madden and Julian, 1994).

4. Cause-effect pathways

4.1. Suspended-sediment

Corals are both autotrophs and heterotrophs and many studies have shown raptorial capabilities and the capture of up to meso/macro sized zooplankton by nematocyst discharges and tentacle grabbing (Ferrier-Pages et al., 2003; Marshall and Orr, 1931; Sebens et al., 1998; Sebens et al., 1996; Vaughan, 1916). Zooplankton feeding contributes significantly to fixed carbon incorporated into coral skeletons (recently reviewed by Houlbreque and Ferrier-Pages (2009)).

From the early suggestions of Goreau et al. (1971) and Muscatine and Porter (1977), corals have been shown to gain some energy hetero-trophically by tentacular suspension feeding (Houlbreque and Ferrier-

Pages, 2009). General suspension feeding mechanisms include direct interception and electrostatic attraction (LaBarbera, 1984; Rubenstein and Koehl, 1977) and once captured there is a second stage involving particle retention and a third phase involving movement of captured particles to the mouth (Shimeta and Koehl, 1997). Tentacular suspension feeding for particulate matter occurs through entrapment in mucus on the coral surface. Another capture process is via mucus threads or filaments attached to the oral disk that are swept by water turbulence entangling fine particulate material as well as larger zooplankton (Lewis and Price, 1975,1976; Lewis, 1976). Ingestion is completed by movement by cilia of particles trapped in the mucus to the mouth (Lewis and Price, 1975; Vaughan, 1916).

Recent studies have shown that the coral cilia can generate vertical flows extending up to 2 mm from the surface which can substantially enhance the transport of solutes to and from the coral surface (Shapiro et al., 2014) suggesting corals may be complementing passive entrapment by active mechanisms. Marshall and Orr (1931) observed that ciliary movement of deposited sediment is a short term response which cannot be maintained for long periods, implying a metabolic cost. However, the energy invested in powering the ciliary movement is reported to be a negligible fraction of the coral's metabolic budget (Shapiro, 2014), but how this changes in response to increased drag associated with high SSCs has yet to be determined.

Ingestion of sediments has been observed in many studies (Lewis and Price, 1975; Lewis, 1976; Logan, 1988; Marshall and Orr, 1931; Stafford-Smith and Ormond, 1992; Stafford-Smith, 1993) and appears


R.Jones etal. / Marine Pollution Bulletin xxx (2015) xxx-xxx

Fig. 4. A conceptual model of the effects of turbidity-generating events on corals based on the US Environmental Protection Agency (USEPA) causal/diagnosis decision information system framework (CADDIS), identifying the sources of sediment, dredging-related turbidity-generating activities, all known cause-effect pathways, biologically plausible cause-effect pathways, modes of action, interacting factors, and likely physiological and ecological responses (see text for explanation).

to be part of a normal feeding mechanism. Suspended particulate matter constitutes a potentially diverse food source containing bacteria, microalgae, protozoa, detrital organic matter (Marshall, 1965), interstitial invertebrates, detached, undissolved mucus (Wild et al., 2004), microbial exudates, and excretory products from other animals (e.g. from fish, Meyer and Schultz, 1985; Lopez & Levinton 1987; Houlbreque and Ferrier-Pages, 2009). Many studies have now shown that after sediment ingestion corals are capable of assimilating and obtaining nutritional benefits from the associated organic matter (Anthony, 1999, 2000; Anthony and Fabricius, 2000; Mills and Sebens, 1997; Mills et al., 2004; Rosenfeld et al., 1999).

Mills and Sebens (1997) suggested that high loads of clean sediments may cause most of the polyps to stop feeding and reject sediments, reducing ingestion rates. Similarly, Anthony (2000) indicated that at concentrations >30 mg L-1 Acropora millepora and Pocillopora damicornis from clear water offshore environments showed a tendency to retract their polyps reducing potential for energy gains from feeding. It follows that depending on organic content, water flow and morphology, low SSCs may be beneficial for some corals in some circumstances and detrimental at higher concentrations. This concept of a low dose stimulation and high dose inhibition has now been reported in numerous studies (Anthony, 1999, 2000; Logan et al., 1994; Mills and Sebens, 1997; Mills et al., 2004; Mills and Sebens, 2004; Rosenfeld et al., 1999; Stafford-Smith and Ormond, 1992; Tomascik and Sander, 1985). Whilst low suspended-sediment concentrations can have beneficial effects,

overall high SSCs is one of the key pressure parameters reducing feeding activity and requiring energy to continually process and transport intercepted sediments (Fig. 4).

4.2. Light availability

Most reef-building corals form mutualistic symbioses with dinofla-gellates of the genus Symbiodinium (Freudenthal, 1962), a diverse range of microalgae divided into nine known clades with significant functional and genetic intercladal diversity (Stat et al., 2012). The algae provide 'photosynthates' or photosynthetically fixed carbon to the host, providing additional energy for respiration and growth (Dubinsky et al., 1984; Falkowski et al., 1984; Goreau, 1959; Lesser, 2004; Muscatine, 1990). The Symbiodinium spp. reside endosymbiotically in the coral endodermal (gastrodermal) tissues within a membrane complex, the symbiosome (Roth et al., 1988; Wakefield and Kempf, 2001), at densities of typically one to two, but sometimes up to six per host cell (Muscatine et al., 1998). Forming the symbiosis enhances deposition of the calcium carbonate skeleton in light-enhanced (DCMU-sensitive) calcification (Gattuso et al., 1999). The highly reflective skeleton also enhances the light field experienced by a polyp through light scattering and diffuse reflection, increasing the probability of absorption and increasing the exposure to photosynthetically active radiation (PAR) by 3-20 fold (Enriquez et al., 2005; Kuhl et al., 1995; Marcelino et al., 2013; Reef et al., 2009).


6 R. Jones et al / Marine Pollution Bulletin xxx (2015) xxx-xxx

Corals and their algal symbionts are superbly adapted to living in both low light and high light environments, exhibiting behavioural morphological and physiological plasticity to maximise light utilization and minimize damage (Roth, 2014). For the host (animal) these include changes in polyp retraction (Levy et al., 2003) and micro- and macro-scale growth morphology (Barnes, 1973). For the algal symbionts this includes changes in optical cross sectional areas and light harvesting capabilities through changes in photosynthetic and accessory pigment concentrations (Anthony and Hoegh-Guldberg, 2003; Falkowski and Dubinsky, 1981; Falkowski etal., 1984; Mass et al., 2010).

The quantity and quality (spectral composition) of the submarine light field is fundamentally important for the physiology and ecology of the coral-algal symbiosis and light attenuation, mediated by absorption and scattering of light by suspended particles (i.e. a step in the causal pathway), is one of the key proximal stressors in the short and especially long-term. In very low or zero light conditions, corals can enter a state of hypoxia and then anoxia; the mode of action is through reduced autotrophy and hypoxia (considered further below).

43. Sediment covering

The immediate response of corals to deposition of sediments on their surfaces is an attempt to self-clean by moving sediments to edges where they are dropped off the colony (Marshall and Orr, 1931). The principal sediment rejection mechanisms identified for a range of coral species, representing different families and a range of growth forms and corallite morphologies are: ciliary action, hydrostatic inflation, tentacle movement, contractions, and mucus entrapment (Bak and Elgershuizen, 1976; Hubbard and Pocock, 1972; Logan, 1988; Marshall and Orr, 1931; Schuhmacher, 1977; Stafford-Smith and Ormond, 1992; Stafford-Smith, 1993; Vaughan, 1916; Yonge and Nicholls, 1931). These are usually referred to as active processes (requiring energy) and most corals employ ciliary action and hydrostatic inflation (sometimes in conjunction with pulsed contractions and polyp expansion), mucus entanglement, and tentacle movement to remove sediments.

A number of other mechanisms have been proposed including movement of sediment by mesenterial filaments, capture of sediment by nematocysts, and sediment ingestion. Commensal crabs become more active during high sediment deposition rates (Stafford-Smith and Ormond, 1992) and can create local turbulence, which dislodges particles. Turbulence from feeding fish can have similar effects (Loya, 1976). It is unclear whether some of these proposed mechanisms are associated with feeding responses and whether they represent physiologically significant sediment-rejection mechanisms. Observations of tentacle movement and processing of individual sediment particles may be a consequence of the large, sand-sized particles used in some of these feeding experiments (Stafford-Smith and Ormond, 1992). Ingested sediments are subsequently regurgitated several hours later as mucous-bound pseudo faeces (Logan, 1988), so sediment ingestion will not clear sediment from the surface.

Corals also have gross and fine scale skeletal morphologies which can assist gravitational forces to remove sediments from their surfaces. These include surface inclination, branch spacing and diameter in arborescent species and aspect ratio and the degree of sphericity in massive species. Finer scale morphology relates to calical size and shape (Hubbard and Pocock, 1972; Lasker, 1980; Logan, 1988; Marshall and Orr, 1931; Vaughan, 1916). Stafford-Smith (1993) also proposed a new parameter, surface smoothness, which is an index of surface microarchitecture and the ability of corals to expand tissues (by the active process of hydrostatic inflation) above skeletal projections, resulting in a shape which better sheds sediments passively. Calix inclination is a key parameter Logan (1988).

Once the sediment clearance rates have been exceeded, sediments will inevitability build-up on a coral's surface and it becomes progressively buried in a sediment deposit (i.e. smothering, Fig. 3F). The

ultimate fate of the underlying tissues is partial mortality (lesion formation), unless the layer is removed by a storm. One of the significant issues associated with smothering is tissue hypoxia, brought about by either light attenuation caused by a sediment covering or a reduction in gas (solute) transfer across diffusive boundary layers (DBLs). Corals are oxygen conformers, routinely experiencing pronounced diel changes in tissue oxygen concentrations ranging from super-saturation (hyperoxia) during the daytime associated with algal photosynthesis, to night-time oxygen shortage (hypoxia) or even anoxia by host and algal respiration (Jones and Hoegh-Guldberg, 2001; Kuhl et al., 1995; Shashar et al., 1996). In darkness, or low flow conditions, oxygen concentrations can fall to levels where aerobic respiration and ATP generation is limited, a situation analogous to flooding of terrestrial plants (Bailey-Serres and Voesenek, 2008; Fukao and Bailey-Serres, 2004). Sea-anemones can survive hypoxic periods by engaging in fermentation processes involving glycolysis (Ellington, 1977; Ellington, 1980,1982) which generates some ATP although at much lower yields (6x less) than aerobic respiration (Shick, 1991). Fermentation processes have been regularly implicated in corals as a means of tolerating short-term hypoxia (Weber et al., 2012; Wooldridge, 2013).

The mechanism underlying sediment-smothering induced mortality and local necrosis has been described in detail by Weber et al. (2012). A rapid (<24 h) microbially-mediated anoxia and change in pH was recorded in Montipora peltiformis smothered with a couple of millimetres of organically-rich sediments, leading to localised necrosis. Sediment smothering is therefore a key pressure parameter associated with turbidity generation, resulting in boundary-related effects and decreased solutes (such as oxygen) and metabolite exchange, mass transport limitations, and decreased filtering/feeding. Sediment smothering is related to sedimentation processes in the water column, which is dependent on wave movement turbulence and local hydrodynamics (i.e. an interacting factor).

4.4. Combined effects of proximal stressors

The key pressure parameters identified above and associated with the indirect effects of dredging activities on reef communities (represented by the inner triangle in Fig. 4) are: (1) high SSCs affecting feeding (heterotrophy reduction), (2) a reduction in light and associated effects on the photosynthesis of the symbiotic microalgae of corals (phototrophy reduction), and (3) sediment smothering which causes a reduction in gas (solute) transfer across diffusive boundary layers. These three cause-effect pathways are highly interconnected, with suspended-sediments causing biological effects directly, but also acting as a causal step to changes in light quality and quantity (through attenuation and scattering in the water column). Similarly, high SSCs are a prerequisite for sediment deposition, mediated by the process of sedimentation (a casual step) in the water column. Once the coral's surface is veneered or smothered by sediment, the effect will be similar to light reduction in the water column and feeding processes will also be affected, and for this reason smothering by sediments has most biological effects associated with it (Fig. 4).

The relative influence of these three key pressure parameters can be symbolized in the ternary diagrams of Fig. 5 with epibenthic filter feeders such a sponges most likely to be more affected by high SSCs (Bell et al., 2015) and sitting at the apex of the triangle (Fig. 5A). Seagrasses, which have comparatively high light requirements (Dennison et al., 1993), are likely to be located more towards the lower left hand side. Non-photosynthetic benthic filter feeders, such as tropical barnacles (Fabricius and Wolanski, 2000), are likely to be located on the right hand-side. Species with mixed modes of nutrition, such as symbiotic hard and soft corals, are likely to be equally influenced and are represented at the triangle's orthocentre (Fig. 5A).

During dredging, these representations are likely to move on an hourly, daily, and seasonal basis, depending and on the dredging activities, diel and tidal cycles, and sea-state. Four different exposure


R.Jones et al./ Marine Pollution Bulletin xxx (2015) xxx-xxx

25 sponge 75

coral barnade



• seagrass LA 25 50 75 SC

where: SS = suspended sediment LA = light availability SC = sediment covering

Fig. 5. Ternary diagrams representing the relative influence of 3 key proximal stressors associated with turbidity generation activities i.e. suspended sediment (SS), light availability (LA) and sediment covering (SC) on shallow tropical benthic organisms such as corals, sponges, seagrass and filter feeders such as barnacles. During dredging the relative influence of the proximal stressors could move on an hourly, daily, and seasonal basis, depending on the dredging activities, diel and tidal cycles, and sea-state. Four different exposure scenarios for symbiotic corals exposed to dredging plumes are symbolized: (B) A scenario whereby a buoyant plume drifts over the reef with little contact with the corals and where light availability only is likely to be the key proximal stressor (see photograph Fig. 3D), (C) A turbidity event during relatively turbulent water conditions where (wave + current) shear stresses are sufficient to inhibit the deposition of most sediment and so suspended-sediments and reduced light availability are the predominant influences (see photograph Fig. 3E). (D) A scenario where elevated SSCs has occurred during very calm conditions and where sediment has subsequently fallen out of suspension and smothered corals (see photograph Fig. 3F), (E) represents a night time scenario of high SSCs in turbulent conditions where suspended-sediment is the sole proximal stressor.

scenarios for symbiotic corals exposed to dredging plumes are represented in Fig. 5B-E. In a scenario of a buoyant plume drifting over the reef with little contact with the corals, light availability is likely to be the key stressor (Fig. 5B and photograph Fig. 3D). Under energetic water conditions, where most sediment is in suspension, suspended-sediments and reduced light availability are the predominant influences (Fig. 5C and photograph Fig. 3E). Fig. 5D represents a scenario where elevated SSCs occurred during calm conditions and where sediment has fallen out of suspension and smothering has occurred (Fig. 3F). Fig. 5E represents a scenario of high SSCs in turbulent conditions at night

time, where suspended-sediments are the predominant influence while light attenuation and reduced light availability has no influence.

5. Water quality characteristics of dredging plumes

There have recently been several major dredging programmes in Australia where the state and federal regulatory conditions have required detailed water quality monitoring programmes over extended periods using arrays of in situ instrumentation (Fig. 6, Falkenberg and Styan, 2014). These data have been made available for study and are

Fig. 6. Location map showing the Pilbara coast in Western Australia and the sites of 3 recent large scale capital dredging projects, including: (A) the Barrow Island project (-7.6 Mm3 of sediment dredged over 530 days from 19 May 2010 to 31 October 2011, Ministerial statement 800), (B) the Burrup Peninsula project in the Dampier Archipelago (-12.5 Mm3 of sediment dredged over 911 days from 22 Nov 2007 to 21 May 2010, Ministerial statement 757), and (C) the Cape Lambert Dredging Project (-14 Mm3 of sediment dredged over 633 days from 22 Dec 2007 to 15 Sept 2012, Ministerial statement 840). These projects were associated with dredging for entrance channels, approach areas, turning basins (TB), material offloading facilities, berth pockets, and liquefied natural gas (LNG) jetties and tug harbours and involved deepening the seabed to —12.5 to — 20 m to allow access for bulk iron ore and LNG transport vessels. The dredging campaigns generally occurred on a 7 days x 24 h basis, stopping only for maintenance and bunkering requirements, with most projects using a combination of trailing suction hopper dredges, cutter suction dredges and back hoe dredges, and bed levellers, and with dredge material placement at offshore disposal sites. The ministerial approval statements (MS) for these projects are searchable on the WA EPA website (


R. Jones et al. / Marine Pollution Bulletin xxx (2015) xxx-xxx

examined below in terms of defining environmentally relevant or realistic conditions during dredging programmes.

5.1. Particle size distributions (PSDs) associated with dredging

Sediment PSDs in the water column were measured during the Burrup Peninsula dredging programme in Western Australia (Fig. 6). Samples were collected inside a plume 125-200 m away from an operating trailing suction hopper dredge and at a reference site outside of the plume and ~800 m southwest of the dredge (Fig. 6B insert). Sampling inside the plume occurred over a 65 min period and at the reference site outside of the plume a further 45 min later. Triplicate water samples were collected at the top, middle and bottom of the water column using a 2.5 L Niskin bottle. Samples were filtered onto Whatman 47 mm GF/F filters, and 100 mLof distilled water used to rinse the container, filter funnel and filter pads of salts. Filters were then dried overnight in a 65 °C oven and weighed. Water column PSDs were assessed using a Laser In-Situ Scattering and Transmissometry (LISST) 100X Type C (Sequoia, WA, US) calculated by examining the angular distribution of forward scattered light (referenced to pure water) over the range from 2-500 |m using the proprietary inversion process. Data shallower than 0.3 m were discarded to remove the possibility of artefacts from small bubbles and dissolving salts from the detector window upon immersion.

Gravimetrically determined depth-averaged SSC ranged from 12-110 mg L-1 within the plume, with the highest SSCs recorded at the surface and the seabed (Fig. 7A) and with depth averaged PSDs ranging within the silt-sized (4-62.5 |m) fraction. Surface plumes had comparatively finer PSDs < 62.5 |m, but peak values of ~60 |m were observed in bottom samples, collected 1 m from the seabed (Fig. 7A). PSDs at the reference site were fine- and medium-sized silts (Fig. 7A). Similar measurements at the offshore spoil ground showed lower SSCs and much smaller particle sizes, with peaks between 10-20 |m and a pattern of slightly coarser sediments at the seabed (Fig. 7C). PSDs of the seabed collected before the start of the dredging programme (see below) indicated a mixture of sand (22%), silt (38%), and clay (40%) was being dredged at the time of sampling.

Grain size analysis of surficial sediments within the area influenced by dredging plumes was examined a few months before and after both the Burrup Peninsula and Cape Lambert projects (Fig. 6). For the Burrup Peninsula project, surficial (top 10 cm) sediment samples were collected by SCUBA divers at ~50 sites along 8 transects with 4 nearshore transects (0.5-2 km from shore) and 4 offshore transects (2-4 km from shore) ranging from 0.1-1 km from the edge of the dredged channel or turning basin (Fig. 6B). Sediments were analysed by a commercial laboratory using Australian Standard (AS) 1289.3.6.2 and 1289.3.6.3 and data expressed as relative percentage of particle sizes in each of the four classes: gravel (>2000 |m), sand (2000-62.5 |m), silt (6.25-2 |m), and clay (<2 |m). For the Cape Lambert project, 5 surficial (top 10 cm) sediment cores were collected by SCUBA divers at ~20 sites along 4 transects (Fig. 6C), with 5 sites along each transect located at distances of approximately 0.25-5 km from the edge of the turning basin (Fig. 6). Sediments were analysed by a commercial laboratory using wet sieving techniques for samples >500 |m and laser diffraction (Malvern Instruments Mastersizer MS2000) for fractions between 2 and 500 |m (ISO 13320-1) and data expressed as relative percentage of particle sizes in each of the four classes: gravel (>2000 |m), sand (2000-62.5 |m), silt (62.5-4 |m), and clay (<4 |m).

The pre-dredging surveys in the Burrup Peninsula dredging project indicated difference in the sediment composition according to proximity to the coast, with the nearshore samples (located within ~2 km of the land) composed of approximately equal fractions of sand, silt, and clay whereas the more offshore sediments (>3 km) were overwhelmingly dominated by sand (~70%) (Fig. 8A). Overall, there are clear differences in the sediment composition after the dredging programme, with

SSC (mg L"1)

1 = 87 2 = 12

5 2 4 3 1 3 = 97

\ V \ \ \ plume \ 4 = 110

- \ reference ... A Reference 5 = 5

10 30 100 300

(B) B SSC (mg L"1) Plume Top =105 Middle =5 Bottom =120

- reference LV plume Reference Top = 5 Middle =0.5 Bottom = 9.5

3 10 30 100 300

(C) Placement site

SSC (mg L-1)

/C\\ Plume

if S M B Top = 5.4

\ / \ \ \ Middle =1.4

■ . A Bottom = 7.2

3 10 30 100 300 Particle size (pm)

clay (4 pm)

coarse sand \ ° fine silt (8 pm)

(1000 mm) \ o medium silt (31 pm)

\ O coarse silt (62 pm)

v. fine sand (125 pm)

/ medium sand \

I (500 pm) I

\ /1 I fine sand (250 pm)

Fig. 7. Sediment particle size distributions in the water column, collected 150 m away from a working trailing suction hopper dredge in a capital dredging programme, showing the fine nature of the released sediments (Burrup Peninsula Project, see Figs. 3A and 6B). (A) Depth averaged particle size distributions (|jm) and suspended sediment concentrations (mg L-1) inside and outside of a dredging plume. Numbers refer to the sites shown in Fig. 6 B. (B) Average particle size distributions at the top (>0.3 m deep), middle (4-5 m) and bottom (8-9 m or 1 m from seabed) of the water column inside and outside of the plume or (C) at the dredge material placement site. (D) Stylized display of coarse sand through silts to clay based on the Udden-Wentworth US standard classification scale of sediments.

increases in the silt content and decreases in the sand/gravel fraction (indicated as a movement of the larger circular symbols in the ternary diagram of Fig. 8). The second post-dredging survey (conducted 6 months after the first) showed a near identical pattern of sediment composition (data not shown). In the Cape Lambert dredging programme, all sediments were sandy (70%) before dredging, with silts and clays making up only 10% and 5% respectively, of the size fractions. After the 2-year dredging programme, there was a reduction in the gravel and sand fraction of the sediment and a near doubling of the silt and clay fraction to 20% and 10%, respectively (Fig. 8B).


R.Jones etal. / Marine Pollution Bulletin xxx (2015) xxx-xxx

Fig. 8. Particle size distributions in surficial sediment before and after dredging, showing a shift to finer grain sizes after dredging. (A) Cape Lambert and (B) Burrup Peninsula dredging projects (see Fig. 6B, C). The accompanying ternary diagrams show clay-silt-sand-gravel size distributions based on the Udden-Wentworth standard classification scale with the larger circles representing the average PSD before and after dredging.

5.2. Suspended-sediment concentrations (SSCs)

At the Barrow Island project (Fig. 6A), turbidity and light levels were recorded on a single sensor platform attached to a steel frame mounted ~40 cm from the seabed. Turbidity was measured using a

single sideways mounted optical backscatter device (nephelometer) and Photosynthetically Active Radiation (PAR) was recorded using a 2n quantum sensor (see Jones et al., 2015a, 2015b for water quality measurements and site descriptions). Data were recorded every 10 min before and during the ~1.5 year dredging programme at

Fig. 9. Turbidity, light and water depth during the Barrow Island project close to the dredging and at a distantly located reference site, showing loss of all benthic light during short-term, dredging related turbidity events. Shown are the NTU and instantaneous PAR (|jmol photons m-2 s-1, primary y-axis) and water depth (metres, secondary y-axis) at Barrow Island at (A) site 1 in Fig. 6A and at (B) a reference site near the Montebello Islands (site 10 in Fig. 6) over a 4 day period in April 2011. Black bars represent night time periods and arrows represent daytime darkness periods when elevated SSC levels have reduced PAR levels to 0 |mol photons m-2 s-1. Turbidity at the reference site is barely detectable, averaging < 1.5 NTU over the 3 day study period.


R. Jones et al. / Marine Pollution Bulletin xxx (2015) xxx-xxx

multiple sites, but for the purpose of this analysis, data were averaged from 3 of the sites closest to the dredging (see Fig. 6A) located 250 m from where the dredging was occurring.

A characteristic feature of the water quality data is pronounced temporal variability, shown in Fig. 9A for a 3-d period in April 2011 where NTUs underwent short term (several hours) 20-fold excursions, from < 5 > 80 NTUs. These events were associated with tidal cycles and turbid plumes passing over the sensor platforms on an incoming tide. When the turbidity levels peaked, the benthic light levels decreased to 0 |mol photons m-2 s-1 during the daytime (see arrows in Fig. 9A). At a reference location with similar water depth (8-10 m) but located 35 km from the dredging and uninfluenced by sediment plumes, turbidity levels were very low, averaging < 1.5 NTU over the 3 day period and mid-day light levels peaked at ~100-200 |mol photons m-2 s-1. This indicates that the complete loss of daytime light near the dredging was caused by turbidity rather than high cloud cover.

Over the 1.5-year dredging programme turbidity levels regularly exceeded 100 NTUs compared to baseline levels of <20 NTUs, but these episodic periods of poor seawater quality were interspersed with periods of good seawater quality (Fig. 10). Given this characteristic, a running means analysis of the raw 10 min turbidity and light measurement was conducted over multiple incrementing time periods from 1 h (for turbidity) or 1 day (for PAR) to 30 d. Each running time period calculated the average of the previous NT data points, where NT is the number of samples in the T hour mean. For example, for the 2-h running mean (T = 2), NT = 12 as there are six 10-min samples per hour. The T hour running mean at a point in time t

Xt (t)= ^ E*i(t) (1)

NT i=1

Where xT (t) is the mean calculated over the previous T hours of the data from time t-Tto time t hours, and xi(t) arethe NTdata points up to and including time t. To avoid biased averages, no xT value was recorded

if more than 20% of the data points for any particular running mean time period calculation were missing. Percentile values of the running mean values xT(t) for each running mean period were then calculated for the pre-dredging and dredging periods.

In R (R Core Team, 2014), running means were calculated by converting the data series for each site into an S3 time series object using the zoo function from the zoo library (Zeileis and Grothendieck, 2005) then applying the runmean function from the caTools library (Tuszynski, 2013). Once running means for each time span were calculated, these were summarized using an average along with various per-centile (P) values (P50, P80, P99 and maximum for turbidity and P50, P20, P5, Pi and minimum for PAR). These were plotted as a function of the running mean time span and compared for the pre-dredging and dredging periods. Analysis of the baseline data (i.e. pre-dredging) captures short term, transient increases in turbidity associated with natural turbidity events, whilst analysis when dredging was underway shows the effect of turbidity generating effects on top of the natural patterns.

Prior to dredging the P95 percentile of the 1 h, 1 day, and 1 week turbidity values were all 2-3 NTUs, whilst during dredging these values were over an order of magnitude higher at 22,18, and 16 NTUs, respectively (Fig. 10A). The 5th percentile of the daily light integral (DLI, mol photons m-2) for 1 day and 1 week running mean periods was ~0.9 and 1.3 mol photons m-2 in the baseline period but 0.1 and 0.3 mol photons m2 when dredging was underway (Fig. 10B). During the dredging programme one of the worst sequences of light reduction was a 16-d period in the first few weeks of May 2011, where the average DLI did not exceed 0.1 mol photons m-2 and the maximum instantaneous PAR was 30 |mol photons m-2 s-1.

Daily light integrals averaged 3.3 mol photons m-2 in the baseline period as compared to 1.8 mol photons m-2 day-1 during dredging. NTU data can be converted to SSCs (as mg L-1) by applying site-specific algorithms (conversion factors) based on gravimetrically determined total suspended solid levels versus nephelometer readings. For the Barrow Island project, the conversion factor calculated this way

Month Time interval (days)

Fig. 10. Changes in turbidity and light before and during a major capital dredging programme where (A) Mean instantaneous NTU (every 10 min) and (B) Daily light integrals (DLI, mol m- 2) during the Barrow Island project at 3 sites closest to the dredging (see Fig. 6A) in the pre-dredging (baseline) or during the dredging phase. Data are the mean of three sites close to the dredging (7-9 m deep and <100-500 m away). Figures on the right hand side show the different percentiles over different running mean periods (from 1 h to 30 days) before (dashed lines) and during (solid lines) the dredging programme.


R.Jones et al./ Marine Pollution Bulletin xxx (2015) xxx-xxx

was 1.3-1.6 and average nephelometrically-derived SSCs over the baseline period were 2.3 mg L-1 (marginally above the ~1.5 mg L-1 precision of the instruments) and 7.1 mg L-1 over the dredging project. The running means/percentile analysis was also conducted for the baseline and dredging seawater quality monitoring sites located from 0.2-15 km away from the principle dredging activity (the creation of the turning basins) during the Barrow Island project at sites for both the baseline and dredging periods and expressed as the P95 (Fig. 11). The turbidity patterns show a rapid initial decay in turbidity with increasing distance and effects on seawater quality could be observed up to 15-20 km from the dredging.

5.3. Light attenuation and light quality

Approximately 80 vertically-resolved downwelling planar irradi-ance light profiles were measured during the Cape Lambert dredging project using a Hydrorad-2 (Hydro-Optics, Biology, & Instrumentation Laboratories, USA). This radiometer provided irradiance measurement values at sub-nanometre spectral spacing. For each vertical profile the irradiance just-below surface Ed (0,\) and the light attenuation coefficient Kd,\ were determined using the BeerLambert law. A general linear relationship between Kd,\ and surface SSC measurements was established for the dredge plumes encountered during sampling. This linear relationship, applied to all wavelengths between 350 and 850 nm, yielded a mass specific spectral attenuation coefficient for SSC, and an offset closely resembling the attenuation spectrum of seawater itself. Additionally, a relationship between clear sky extrapolated just-below surface incident irradiance Ed (0, \) spectra and the solar zenith angle was determined.

Using these empirical relationships a downwelling irradiance spectrum was simulated for a given sun angle and water column averaged SSC concentration and PAR values were calculated from the full-spectrum measurement. Although empirically tied to the conditions found during the Cape Lambert dredging campaign, the model outputs demonstrated the general and spectral trends encountered in a dredge plume due to increased SSCs (Fig. 12A, B). This included a decrease in light with depth and increasing SSCs and a shift to more yellow light at high SSCs. For reference purposes, a vertically integrated 10 mg L- 1 suspension of silt-sized sediments reduced light levels to ~11 ^mol photons m-2 s-1 at 10 m depth and a 30 mg L-1 SSC reduced the light to the same values by ~4 m depth (Fig. 12A). At 5 m depth, the light spectrum under a dredging plume shifted almost entirely to yellow/green wavelengths at

concentrations of 10 mg L-1 and higher (Fig. 12B). These calculations were simulated for a clear (cloud-free) sky at solar noon, and during cloudy days at lower azimuth angles and at different sea states, underwater light quality and quantity would be substantially lower.

6. Experimental studies of the effect of sediments on corals

Laboratory-based studies examining the effects of sediments on corals can be grouped into experiments examining the effects in suspension and those examining the response of corals to a downward flux of particles. The latter group includes a range of burial and sediment covering experiments. As discussed further below, it is not clear if sediments were kept in suspension in the former group and the division between the tables is subjective.

6.1. Suspended-sediments

Some of the earliest studies of the effects of suspended-sediments were associated with understanding the sub-lethal and lethal toxic-ity of drilling muds and fluids on corals (Kendall et al., 1983; Szmant-Froelich et al., 1981; Thompson, 1980; Thompson et al., 1980) and several short-term feeding experiments investigating the effects of particulate matter concentrations on ingestion of corals (Table 1). However, the majority of the studies have been conducted over longer terms (weeks to months) and associated with examining the effects of sediments on corals to gain a better understand the effects of river runoff and/or dredging (Table 1). Whilst some studies have been conducted in situ using sediments introduced to small chambers held on the reef (i.e. Thompson et al. (1980) and Kendall et al. (1983)), most experiments were conducted in aquarium systems associated with marine research facilities. These ex situ studies have been conducted using either artificial lights (fluorescent and metal ha-lides lamps) or under natural sunlight (<300 ^mol photons m-2 s-1) using neutral density shade cloth to manipulate irradiance.

Sediments used in these studies have been calcium carbonate sands from the reefs where the corals were collected, sediments collected by filtering or back flushing filters, marl (a naturally occurring calcite), and kaolin clay (Table 1). The particle size distributions in these studies have not always been quantified and range from fine sands (i.e. Rice and Hunter 1992; Sofonia and Anthony, 2008), to fine silts (<20 |jm, Flores etal. (2012). Acombination of water pumps and aeration have typically been used to keep sediments in suspension.

• 95th Percentile

f 2 • Dredaina period

Fig. 11. Turbidity (NTU) values at different distances from dredging (from 200 m to 15 km away) and over different running mean time periods (from 1 h to 30 days). Shown are the 95th percentile values (see (1)) in the baseline period (before dredging, grey symbols) and during the Barrow Island dredging project (black symbols, see Fig. 6A).


R. Jones et al. / Marine Pollution Bulletin xxx (2015) xxx-xxx

Fig. 12. Simulated effects of suspended sediment on light quantity and quality based on empirical data collected during the Cape Lambert dredging project. (A) Simulated vertical PAR (|mol photons m-2 s-1) profiles for eight SSCs from 0.5-30 mg L-1 (B and C) Simulated downwelling irradiance spectra at 5 and 10 m depth with varying SSCs from 0.5-30 mg L-1 and associated PAR as |mol photons m-2 s-1. The incident downwelling irradiance spectrum for all figures is typical of a clear sky at tropical noon, with an initial (underwater, 0 m depth) PAR value of~1530 | mol photons m-2 s-1.

6.2. Sediment deposition

The sediments used in deposition studies have typically been collected from local reefs or inter-reefal area and primarily composed of biogenic calcium carbonate or more terrestrial, siliciclastic sediments collected from river mouths (i.e. Loiola et al., 2013) (Table 2). One study used terrestrial quartz/granite beach sand (Peters and Pilson,

1985) and several studies have used silicon carbide (carborundum) for experiments on smaller particle sizes (e.g. Bak and Elgershuizen, 1976; Stafford-Smith, 1993; Browne et al., 2014) and limestone (Logan, 1988). Most studies employed a screening process to remove coarse and fine material and although PSDs of the sediments have only been quantified in a few studies, most have used sands, a few have used silts, and only the more recent studies have begun using fine, silt sized fractions, including the studies by Weber et al. (2006), Flores et al. (2012) and Weber et al. (2012).

The application methods have varied widely from manual application of sediments via a tube (i.e. Bak and Elgershuizen, 1976), funnel (Riegl, 1995), or pipette (Logan, 1988), a syringe (Gleason, 1998) or manually applied by unspecified techniques (Schuhmacher, 1977; Lasker, 1980; Stafford-Smith, 1993; Sofonia and Anthony, 2008; Piniak, 2007). In some of the studies, sediments were applied by creating initially high SSCs, which were then allowed to settle out of suspension on the corals (Philipp and Fabricius, 2003; Weber et al., 2012; Weber et al., 2006). For longer term exposures various sediment resuspension apparatus have used re-circulatory airlift systems (Browne et al., 2014; Riegl and Branch, 1995; Todd et al., 2004).

The majority of studies reported nominal sedimentation rates calculated by weight of sediment added and the surface area of the colonies. In some studies, it is not clear how sedimentation rates were calculated (Lirman et al., 2008) and in a few studies, sedimentation rate was measured using small sediment traps or plastic squares placed within the experimental containers (Browne et al., 2015; Flores et al., 2012; Peters and Pilson, 1985; Sofonia and Anthony, 2008; Todd et al., 2004).

7. Discussion

Our present day understanding of the effects of sediments on corals comes from observations and the results of many laboratory- and field-based manipulative studies in a body of literature generated primarily over the last 30 years. It has been reviewed many times (Erftemeijer et al., 2012; Fabricius, 2005; Foster et al., 2010; Jones et al., 2015b; Rogers, 1990), and it seems reasonable to assume that results from these studies can be used for impact prediction purposes and deriving water quality values for managing dredging programmes near coral reefs. However, it is noticeable that seawater quality conditions during dredging programmes have never been quantified in detail (but see Jones et al., 2015a), and using information from several of the large scale dredging programmes close to coral reefs in recent years, the in situ seawater quality conditions differ in many regards from the experimental conditions used in past manipulative studies. In their review of principles of sound ecotoxicology and risk assessment, Harris et al. (2014) emphasize the critical importance of defining and testing realistic and environmentally relevant exposure scenarios and to comprehensively justify those exposure conditions (Harris et al., 2014).

The conceptual model developed in this review identified key cause-effect pathways affecting corals are light attenuation affecting photosynthesis (autotrophy), high SSCs affecting feeding and cleaning processes (heterotrophy), and sediment covering restricting solute exchange (smothering). These are well-known cause-effect pathways for the effects of sediments on corals (see Rogers, 1990), but from the conceptual model it is clear the proximal stressors are highly interlinked, with some connected along causal pathways and some acting alone or in combination. As such, the most relevant parameter(s) may change according to dredging activities, sea-state, distance from the dredge etc. Homer's sea-god Proteus was known for his ability to transform, i.e. to change shape and form. The 'protean' nature of suspended-sediments makes it difficult to identify which is the most relevant or important pressure parameter(s) at any given time, to identify cause-effect pathway(s), and importantly, to establish dose-response relationships. For example, if smothering of the coral's surface occurs, any biological effects which subsequently occur cannot be related to turbidity measured in the overlying water column. This has important

.O m CL. DJ

1 s O a

a : p j

°o O ,

2 R o ,

Studies examining the effects of suspended-sediments on corals. Experiments were conducted either under diel cycles or artificial lights from fluorescent and or metal halide lights. Under artificial lighting, the light intensities are expressed in |jmol photons m2 s-1 and Daily Light Integral (mol photon m day-1) calculated from the photoperiod. ns = not specified, SPM = suspended particulate matter.


Sediment type

PSD (Mm)

Test type

Time (day)

SSCs (mg L-1)

L:D cycle Light source

PAR (max |jmol m 2 s 1 ) DLI (mol m-2 day-1)

cre o 3 c

PAR 2-2.5, DLI >0.13

Rice and Hunter , . •>•>•.. ™ 4conc. 14:10 h

,„„„_, 7 species of Atlantic corals Offshore reef sediments <500 33 min turnover 10-20 „„ „„„

(1992) 49-199 Source: ns

Experimental outcome: No effects on survival in any treatments and no consistent patterns of a decline in growth rate

1-2 NTUs: PAR-68, DLI 3.4 7-9 NTUs: PAR-64, DLI 3.2 14-16 NTUs: PAR-62, DLI 3.1 28-30 NTUs: PAR -56, DLI 2.8

Experimental outcome: Effects on P/R ratios at 14-16 and 28-30 NTUs compared to controls. Prolonged exposure to 28-30 NTUs caused lesions in both species. NTU of 29-30 NTUs was equivalent to a Marl concentration on 55 mg L-1.

Telesnicki and Goldberg (1995)

D. stokesii M. meandrites

Silt sized marlstone

Partial water changes every 4 days

1-2, 7-9,14-16, 28-30 NTUs

14:10 h

Metal halide lamps

Anthony and Fabricius (2000)

G. retiformis P. cylindrica

Collected by back-flushing a sand filter

33 min turnover


Filtering of water from the reef

Experimental outcome: Skeletal growth rates negatively affected by shading but not by the SPM treatments

P. damicornis A. millepora

Experimental outcome: Experimental outcome: Reduced feeding rates in corals exposed to 30 mg L-1

Sofonia and Anthony Collected from an inshore

Turbinaria mesenterina (2008) turbid reef

Experimental outcome: No effect of the highest sediment load, regardless of flow conditions



Flow though with repeat dosing

Flooes et al. (2012)

A. millepora M. aequituberculata

Reef sediment ground using a ceramic mill

Mean 6 1.3 day turnover

<2 -4 -16

1,4,8,16 and 30

<1,16,100 mg cm2

1,3,10, 30,100

Natural cycle

Sunlight + neutral density shade cloth

Natural cycle

Sunlight + neutral density shade cloth

10:14 h

Metal halide lamps 12:12 h

Fluorescent lights

<2: PAR 600, DLI 8.1-12.8 -4: PAR 140, DLI 2.5-4 -16: PAR 140, DLI 2.5-4

PAR equivalent to 3-5 m depth across all treatments

PAR-300, DLI 11

1mgL-1: PAR 200, DLI 8.3 30 mg L-1: PAR 184, DLI 8 100 mg L-1: PAR177, DLI 7.6

Experimental outcome: For branching A. millepora some partial mortality occurred in the 100 mg L 1 treatment (LOEC) but no mortality was noted at 30 mg L 1 (NOEC). For M. aequituberculata (horizontal growth form) partial mortality was noted at 3 mg L-1 and above, but related to settlement of sediment on the surface.

Cooper and Fabricius (2012)

Porites sp

Collected from an inshore reef

Experimental outcome no mortality in corals, change in colour brightness in corals exposed to SPM

Browne et al. (2015)

Merulina ampliata, Pachyseris speciosa, Platygyra sinensis

Silicon carbide

Mean 60

Continuous flow

Flow through

2 and 20 under 2 shade treatments

50-100 100-250

12:12 h

Metal halide lamps + neutral density shade cloth

10:14 h

High Output aquarium bulbs

1-2 mg L-1: 835, DLI 37 20 mg L-1: PAR 598, DLI 26 1-2 mg L-1: PAR 56, DLI 2.4 20 mg L-1: PAR 32, DLI 1.4

1 mgL-1: PAR140, DLI 5 50-100 mg L-1: PAR 110, DLI 4 100-250 mg L-1: PAR 80, DLI 2.9

Table 2

Studies examining the effects of settling sediments on corals.



Bak and Elgershuizen (1976)

19 Caribbean reef corals

Reef sand and carborundum powder

Schuhmacher (1974) 14 species from Gulf of Aqaba, Red Sea Carbonate and silicate sand


Moritastrea cavernosa

Collected from a reef, wet sieved and 2 fractions used

Peters and Pilson (1985) Astrangia danae

Quartz/granite beach sand passed through <500 |jm sieve and combusted at 550 °C

Logan (1988)

Scolymia cubensis

Fine, medium and coarse

Hodgson (1990)

Oxypora glabra, Montipora verrucosa, Fine, freshwater washed and Pontes lobata, Pocillopora meandrina dried marine sediment

Stafford-Smith (1993) 22 species of Australian corals

-70% carbonate & 30% quartz obtained from reefs

Riegl (1995)

8 species of corals from South Africa Locally collected non-reef sand

Favia favus, Favites pentagona, Riegl and Branch (1995) Platygyra daedalea, Gyrosmilia interrupta

Coarse reef sand combusted at 350 °C

PSDs (|jm)

Application methods

Application rate (mg cm day-1)

Experimental outcome

100-3000 mean = 1200

>180 < 1.4 mm

60-250(fine) 500-1000 (coarse)

62-500 mean = -200


'Fine' sediment

63-250 500-1000

v. fine (40-125) fine (125-250) coarse (>500) sand

46.1% 250-500 53.4% 125-250

0.75,1.5 and 3.5 g sand and 0.75 g carborundum delivered by a tube

Sand dropped on corals to cover the surface by a layer of 1 mm

20 cm3 of sediment deposited on each colony

Sand sprinkled on the surface of the corals which were examined for 8 h

3 g spread with a pipette over the corals' oral disk

50 or 300 g of sediment stirred vigorously 1-2 x day

5 g of sediment spread evenly over a 5 x 5 cm surface of the coral

100-425 (sand)and 100 (carborundum)

Could not be determined

18.5 (fine) 74.4 (coarse)

200 applied daily for 4 weeks then 3 times a day for 2 weeks

Cannot be determined but likely to be >100 s

30-40 for 7-10 day

200 and 200 for 6 weeks

Deposited through a funnel and a constant flow of fine sand through a re-circulatory system

14.2 g applied to a chamber containing a coral

200 for 8 h 200 for 6 weeks (continual coverage)

Sand cleared more efficiently than carborundum powder and coral surfaces were typically clean by 24 h in up to 3000 mg cm2 treatments Most species covered with 1 mm of sand were still partially covered after 72 h

Approximately half to two thirds of the sediment removed from the surfaces within 8 h with most clearance occurring in the first 2 h Sand rapidly (within 1-2 h) cleaned from the surface with little remaining after 24 h. No effects at the 200 mg cm-2 day-1 treatment but some cellular damage at the 600 mg cm-2 day-1 treatment 10-60% removed within 24 h in horizontally orientated corals, with fine sediments removed quicker than coarse sediment. Sediment clearance was 90% complete within 24 h if corals were inclined at 75° Sediments accumulated on flat and concave parts resulting in bleaching then tissue necrosis, and sediments overlying tissues changed to a greyish-black

Fine sediments were rejected more efficiently than coarse ones, with branching species clearing sediment loads within a few hours. Clearance of 50-100% of surface occurred within 24 h, with some less tolerant species failing to completely remove sediments after a few days. Bleaching and mortality was observed in some species within a few days, but other species required longer periods of sediment covering for mortality to occur No difference between sediment type and 50-100% clearance of sediment in 8 h with no differences between species in the ability to shift different grain sizes. Mortality noted in 3 species after 15 days continuous exposure to 200 mg cm-2 day-1

Reduction in P/R ratios at all concentrations

Gleason (1998)

Pontes astreoides

Intertidal beach sand Combusted at 450 °C

Philipp and Fabricius (2003)

Moritipora peltiformis

Collected from 3 m water depth from a local harbour

Todd et al. (2004)

Diploastrea heliopora Favia speciosa

90%: 10% quartz: carbonate

Weber et al. (2006) Moritipora peltiformis

Collected from 2 rivers estuaries and 3 reefs, and aragonite dust from Pontes skeletons

Piniak (2007)

P. lobata andM. capitata

Carbonate beach sand and siliciclastic harbour mud

Sofonia and Anthony (2008)

Turbinaria mesenterina

Sediment (sandy silt) collected from a turbid reef environment

Lirman et al. (2008) Pontes astreoides Siderastrea siderea

Surficial sediments collected from a local reef

Flores et al. (2012)

Acropora millepora Montipora aequituberculata

Sediments were collected from an offshore reef

Weber et al. (2012)

Montipora peltiformis

Organic rich sediment collected from 5-10 m on a fringing reef of GBR

Loiolaet al. (2013)

Mussismilia braziliensis

Muddy, 91% siliciclastic sediment from a river channel

Browne et al. (2015)

Merulina ampliata, Pachyseris speciosa, Platygyra sinensis


Mean = 253 70% <10,

25% 10-50 »,•_,., • 5-6 applied twice a day

3% > 50-100 Applied with a syringe forfourdays

2% >100

Fine, muddy sediment screened to <1000

Unspecified amount of sediment was placed into a 1000 L tanks and allowed to settle over 6 h

Average 151 (Range 79-234)

5% coarse, 45% medium, 45% fine and 5% very fine sand

Unspecified amount of sediment was resuspended for 1-5 min four times per day.

Average of 20 for 4 months

<63, fine sand (63-250) medium (250-500)

Unspecified amount of sediment was placed in 60 L containers and allowed to settle out over a 2 h period

Nominal values of 33,66, 100,133,160

Sand (99%, 63-2000), mud (55%, 63-2000 and 23% < 0.62)


200-250 g of sand or mud spread evenly over the surface of the corals

Fine sediment was applied manually 3 x per day for 34 days

M. capitata = 2200 (mud), 2,800 (sand), P. lobata = 1,500 mg (mud), 1,600 (sand)

<1,16 and 100 each day

Sand with mean PSD of 176

1000 mg L 1 sediment added daily to 5 L containers

53 per day for 3 weeks

Added to 30 L containers, 95% <20 |jm mean size of and kept in suspension by <6|jm pumps to give 1,3,10,30,

100 mg 1 solutions

<63 |jm mean = 15 |jm

Fine silt

1 -300 |jm median 60 |jm

Sediment added to 60 L containers and held static for several hours to coat corals in a layer of sediment

Sediment was added to 4 L containers and stirred for 2 min each day to resuspend the sediments

Corals exposed to intermittent bursts of sedimentation

66 = 2.1 -2.6 mm thick

Unspecified amount of sediment allowed to settle over 24 h to produce 7 sedimentation rates of 0,10,50,199, 250, 350 and 450

4 week exposure to 25-65 and 4 week recovery period

Sediment built up on the corals' surface resulting in many colonies exhibiting

Bleaching and early signs of necrosis in colonies exposed to 151 mg cm~2 for 12-18 h and necrosis in corals covered from 24-36 h. Effects were confined to areas that were sediment covered Some minor bleaching was observed, no mortality occurred and no obvious morphological changes Sands were removed more efficiently than silts and nutrient poor sediments removed more efficiently than nutrient enriched ones. Anoxia develop under nutrient enriched silty sediments even under short term (days) exposure Reduction in quantum yield in corals smothered in sediments for 30 h or more, with harbour mud having a greater effect than beach sand

No effects observed

Reduced growth rate in corals exposed to sand deposition for 3 weeks and involving a temporary covering of sediment and temporary reduction in light of 95% for 6 h each day No mortality in A. millepora exposed to 30 mg 1 for 4 weeks and some partial and whole colony mortality at 100 mg L-1. Partial mortality in M. aeqiiituberailata occurred as to as low as 3 mg 1 and 60% mortality occurred at 100 mg L_1. Mortality was directly related to sediment smothering which varied considerably between morphology the branching and plate like mortality Rapid (15-18 h) microbially mediated anoxia and tissue necrosis in corals covered in a 2-3 mm thick layer of organically enriched sediments

No necrosis in corals exposed to up to 150 mg cm~2 day-1 of organic free sediments for 45 days or up to 400 for 5 days

No effects (Platygyra sinensis) <5% partial mortality (Merulina ampliata) or<15% mortality (.Pachyseris speciosa)


16 R. Jones et al / Marine Pollution Bulletin xxx (2015) xxx-xxx

implications and potentially confuses and confounds laboratory and field experiments and the associated conclusions.

Summarizing the detailed seawater quality information in this study from Figs 7-11 (and also Fisher et al., in press ; Jones et al. 2015a), in the near-field environment sediments released into the water column and moving out of the immediate dredging area (to create the far-field plume) are primarily silt- and clay-sized. Dredging increases the frequency of extreme values and alters the intensity, duration and frequency of the turbidity events over background levels. Upper percentile values (e.g. > P95) of seawater quality parameters can be highly elevated over short periods i.e. increasing by 2-3 orders of magnitude over a day and exceeding hundreds of mg L-1 over a period of hours. Over longer periods (days), SSCs P95 are in the tens of mg L-1 and typically less than 10 mg L-1 over a period of weeks and months. Scattering and light attenuation by the suspended-sediments occurs rapidly in the water column, with the shallow reef environment routinely experiencing semi-dark, caliginous, or 'twilight' periods, and frequently complete loss of light at moderately elevated SSCs. However, a more common feature was extended periods, i.e. days to weeks, of low light. Different wavelengths are preferentially attenuated underneath plumes, with more immediate loss of red and blue light and a shift to less photosyn-thetically useable yellow-green light. The impacts of dredging on sedimentation and the problems associated with measuring sediment deposition and the significance of quantifying sediment deposition for understanding coral mortality in dredging programmes are discussed further below.

7.1. Experimental studies of the effects of suspended-sediment and light attenuation on corals

It is not clear from the description of many of the laboratory based studies whether sediments were truly kept in suspension throughout the study, especially those with larger particle sizes (i.e. Rice and Hunter 1992) and higher sediment concentrations (Browne et al., 2015; Sofonia and Anthony, 2008). Many of the laboratory-based studies examining the effects of suspended-sediment have not specified the particle sizes and therefore it is difficult to ascribe any observed effects to SSCs or to sediment covering. All studies were conducted in shallow containers and it is conspicuous that many studies have not compensated for the fact that light attenuation is exponential, and likely to be quite small in such a shallow water depth even at higher concentrations. Flores et al. (2012) measured a < 10% reduction in PAR in 15 cm deep experimental containers caused by a 30 mg L-1 SSC of fine silt, whereas on a reef at solar noon and on a sunny day, nearly all light would be attenuated by 5 m under an equivalent concentration (Figs. 9,10 and 12). Some studies have recognised this issue and compensated for this depth effect by reducing light levels with neutral density shade-cloth at higher sediment concentrations, or partially compensated for the effect by reducing light uniformly across several different sediment concentrations (Anthony, 2000; Anthony and Fabricius, 2000; Cooper and Fabricius, 2012). Other studies have not addressed the issue at all (Browne et al., 2015; Flores et al., 2012; Hodgson, 1990; Riegl, 1995; Sofonia and Anthony, 2008; Telesnicki and Goldberg, 1995; Thompson et al., 1980). This can make the results misleading for hazard assessment purposes, i.e. corals could be living through SSC treatments in laboratory-based experiments which in situ could have more profound effects because of the associated light reduction. In the experiments of Browne et al. (2015), PAR levels in the shallow containers during high sediment pulses (50-100 and 100-250 mg L-1) were reduced from 140 |mol photons m-2 s-1 to 100 and -70 |mol photons m-2 s-1, respectively. Because there was no additional shading to reduce the light levels to the levels corals would experience in situ under similar suspended-sediment concentrations, the corals lived through the 4 week exposure with either no effects (Plattygyra sinensis) < 5% partial mortality (Merulina ampliata) or < 15% mortality (Pachyseris speciosa). Survival of corals in situ under more appropriate light regimes (i.e.

most probably full light extinction — see Fig. 12A) over a similar extended time period, could have resulted in a different outcome given the physiological effects known to occur in corals in darkness (DeSalvo etal., 2012; Rogers, 1979; Yonge and Nicholls, 1931).

Compensating for the depth effect using neutral density shade cloth over containers will not, however, correct for spectral changes which may occur underneath dredge plumes. The underwater light field is modified by the spectral-dependent absorption and scattering properties of the seawater itself involving absorption of the red-infrared-spectral region, and also by phytoplankton, humic substances (gelbstoff) and especially particulate matter (Jerlov, 1976; Kirk, 1994; Kirk, 1985). The hyperspectral data showed a reduction in blue and red wavelengths and a clear shift to yellow-green conditions underneath a dredge plume. This region is outside of the major absorption peak of photopigments, meaning that light is poorly absorbed by the corals in this part of the spectrum and it is relatively inefficient at driving photosynthesis (Halldal, 1968; Szabo et al., 2014). This is a significant issue with monitoring programmes and for deriving in situ dose-response relationships from laboratory experiments, because the types of light sensors commonly used in monitoring programmes integrate across the PAR wavelengths and do not account for any spectral changes. That is, the reported light levels in 9,10, and 12 could be misleading as to what corals can actually use for photosynthesis, i.e. the difference between photosynthetically useable radiation (PUR) (Morel, 1978) and photosynthetically active radiation (PAR) (Tyler, 1966). A number of recent studies have begun investigating the effects of light quality on coral physiology (Wangpraseurt et al., 2012; Wangpraseurt et al., 2014) made easier by recent advances in lighting technology (Wijgerde et al., 2012; Yeh et al., 2014), and spectral changes should be considered in future studies if results from laboratory based studies examining SSCs and light reduction are to be extrapolated to the field.

The results from those experiments where there was no light compensation have inadvertently provided evidence to suggest corals can actually survive quite high SSCs as long as the light is sufficient and there is no sediment accumulation on the surfaces. For example, there was no mortality in A. millepora exposed to mg L-1 for 30 days (Flores et al., 2012), or in Porites spp. exposed to 20 mg L-1 for 56 days (Cooper and Fabricius, 2012). Skeletal growth of Goniastrea retiformis and Porites cylindrica exposed to 16 mg L- 1 for 2 months were also not different from ex situ or in situ controls (Anthony and Fabricius, 2000). These sediment loads (intensity x duration) are high compared to natural resuspension events (Larcombe et al., 1995) and conditions which can occur close to a major capital dredging project (see Figs. 10, 11). By isolating SSCs as the primary variable, these studies have provided evidence that for adult corals, light availability and sediment covering are probably the most important cause-effect pathways associated with turbidity generating events in the short term.

7.2. Experimental studies of the effects of sediment deposition on corals

There have been many studies examining the effects of sedimentation on corals but there are also some methodological issues which make interpretation of these studies difficult. Many studies have used silicon carbide as a sediment proxy (Browne et al., 2015; Junjie et al., 2014; Lui et al., 2012; Stafford-Smith and Ormond, 1992), which because of differences in specific gravity, sphericity and porosity and light scattering and absorbing properties makes generalization extremely difficult (Storlazzi et al., 2015). The use of black carborundum, which traces back to the feeding experiments of Yonge (1930), should be considered carefully if the intention is to use the results of such studies to make generalizations of the effects of sediments released from dredging. As significantly, it is noticeable that many studies have used sands (62-2000 |m) as opposed to the fine silts and clays which typify the near and far-field dredge plume. Smaller particles have much greater scattering and absorption properties and so light transmission and solute exchange will vary considerably depending on sediment type


R.Jones etal./ Marine Pollution Bulletin xxx (2015) xxx-xxx 17

(and colour) (Storlazzi et al., 2015). Weber et al. (2006) recorded light transmission of only ~0.1% through a 66 mg cm sediment layer on a coral in contrast to Riegl and Branch (1995), who recorded light transmission as high as 30% through a 200 mg cm2 sediment layer. The primary difference between the studies was that Weber et al. (2006) used silt (with a median grain size of ~10 |am) and Riegl and Branch (1995) used much coarser sand with a median grain size of 250 |am (~50% between 250 and 500 |jm).

The difference between the silt and clay sized particles found in dredging plumes and the sands used in some of the laboratory experiments makes it difficult to interpret information and extrapolate to conditions during dredging. For example, in experiments designed to partition the effect of turbidity and settling particles on Galaxea fascicularis and Goniopora somaliensis, Junjie et al. (2014) acknowledged that settled sediment could create an additional barrier to light, but then discounted the potentially confounding effect on the high light availability reported by Riegl and Branch (1995) discussed above. Junjie et al. (2014) used substantially smaller grain size (10-300 |jm with a median grain size of 60 |jm), used black carborundum (see Storlazzi et al., 2015), and the possibility that light attenuation by the sediment layer was not a significant factor in this experiment study has not been discounted.

Why such coarse sediments have so routinely been used is not clear, but perhaps the reason is purely practical, as silts and clay cloud the sea-water so much that the corals cannot be seen (see Bak and Elgershuizen, 1976; Todd et al., 2004). Settling velocities of sediments are related to density and proportional to the diameter squared according to Stoke's law. Assuming particles do not coagulate, flocculate, or grow and in the absence of vertical mixing, a fall velocity of 1.29 cm s-1 for sand-sized (200 |jm) particles (Storlazzi et al., 2011) should result in sediment at the surface reaching a 10 m deep seabed within < 15 min. Settling velocities of silts are lower and they have the opportunity to move away from dredges onto nearby habitats. Far field plumes (km away from dredges) are likely to be made up of smaller particles still, similar to the fine silts and clays which only recently have been reported as dominating river plumes a few km from river mouths (Bainbridge et al., 2012).

A different and perhaps more fundamental problem than sediment type and particle size is the application rates (sedimentation rates) used in these studies. All the studies have used experimental sedimentation rates based on in situ measurements with sediment traps. The problem with traps is that they capture all particles including those which are just passing over the reef or that are only momentarily deposited (Field et al., 2012; Risk and Edinger, 2011; Storlazzi et al., 2011; Thomas and Ridd, 2004). Traps can even collect sediments that never actually settle and therefore can provide a sedimentation estimate (even in net erosional environments), i.e. they provide a 'pseudo-sedimentation' rate. Interpreting data from traps is more difficult in shallow, energetic environments that are characteristic of reefs, and these limitations do not include the effects of waves and wave orbital velocities which result in acceleration and deceleration of horizontally moving particles over the trap mouth during agitated sea states (Bothner et al., 2006; Storlazzi et al., 2011)(cf Fig. 1). Traps have been described as at best only providing a vague approximation (Storlazzi et al., 2011) of the amount and type of sediment that deposits, and have been suggested as being more apt to record information on suspended-sediment dynamics than provide any useful data on sedimentation (Buesseler et al., 2007; Gardner, 1980; Gardner et al., 1983; Storlazzi et al., 2011). These problems have been discussed and reviewed many times in different fields of marine research, see for example (Bothner et al., 2006; Browne et al., 2012; Buesseler et al., 2007; Butman et al., 1986; Jurg, 1996; Kozerski, 1994; Reynolds et al., 1980; Risk and Edinger, 2011; Storlazzi et al., 2011; Thomas and Ridd, 2004).

A number of studies have attempted to overcome the trapping artefact and address the question of what are typical sedimentation rates on

coral reefs. On Kenyan reefs, McClanahan and Obura (1997) noted that a factor of 3 is required to scale up sedimentation rates on flat tiles (which do not suffer resuspension limitation and deposition bias) to those measured with sediment traps (range 3-6 mg cm-2 day-1). More recently, using circular, concrete-filled PVC hubs (SedPods), Field et al. (2012) measured net sedimentation rates of 0.3-0.6 mg-2 day-1 in Hanalei Bay, Hawaii, as opposed to sediment trap accumulation rates of 7-17 mg cm-2 day-1. Using shallow trays, which also allow resuspension, Browne et al. (2012) estimated that net deposition rates in the high-turbidity nearshore coral settings on the inner shelf of the central Great Barrier Reef, averaged 3-7 mg cm-2 day-1 over the course of a year. These values were considerably lower than measured using traps in the same area i.e. a mean of 44 mg cm-2 day-1 with an upper value as high as 364 mg cm-2 day-1 (Mapstone et al., 1992). These more recent studies of deposition rates need to be contrasted with earlier suggestions — but based on the current understanding at the time — that sedimentation rates in excess of 200 mg cm-2 day-1 for periods of days to weeks are '...not uncommon on fringing reefs of the GBR.' (Stafford-Smith, 1993).

The problem with interpreting information from traps led Storlazzi et al. (2011) to conclude that prior research results in the literature need to be interpreted carefully and with recognition that there may be irregularities in the trapping technique or in the application to understanding coral reef processes. Within the context of developing thresholds for dredging projects, if sediment traps are overestimating sedimentation rates (through resuspension limitation and deposition bias), then replicating those rates conditions in laboratory studies may result in exposure scenarios that are unrepresentative of anything but extreme conditions. The experimental application rates are often in the high tens of mg cm- 2 day-1, commonly hundreds of mg cm-2-day-1, and sometimes even g cm-2 day-1. When these rates are applied under still or low flow conditions this can result in sediment deposits that are millimetres thick on the corals' surface. The deposition rateof66mgcm2day-1 in the studies ofWeberetal. (2012) resulted in a smothering of the coral tissues in a 2-3 mm thick layer of sediment, and this is likely to be less than the earlier deposition experiments of Philipp and Fabricius (2003) where deposition rates of up to — 230 mg cm2 day-1 were used. For comparative purposes, the visually significant turbid flood plumes on the Great Barrier Reef have concentrations of sediments which, if settled to the seabed, would produce a deposit of only 10 |am (Orpin et al., 2004). The question then becomes how representative are experiments that create mm thick deposits of sediment in less than a day, and do these reflect conditions that can occur naturally or only very extreme events such as cyclones or only conditions that occur very close to dredging.

Smothering is regularly observed in dredging projects and occurs because the sedimentation rate is likely to exceed any natural rates corals have experienced previously. This is because dredging can cause high SSCs in sea-states where ambient hydrodynamics cannot support the load (Fig. 3A-D) and the sediment 'overburden' rapidly falls out of suspension according to particle specific settling velocities. Smothering occurs when sediment cannot be cleared fast enough from the surface and sediment begins to accumulate over successive days producing the sort of images in Bak, 1978; Foster et al., 2010, and Fig. 3. In contrast, natural high SSCs produced by high wind events occur where wind-driven waves and tidal currents create conditions where wave orbital velocities are sufficient to keep some of the sediments in suspension and deposition subsequently occurs (after a settling lag) during quiescent periods and after further entrainment and dilution (Ogston et al., 2004). These are very different scenarios and sediment deposition is therefore different from other proximal stressors such as elevated SSCs and light reduction, as short term turbidity events and low light, twilight, periods are not uncommon in the marine environment associated with wind and wave events (Anthony and Larcombe, 2000; Jones, 2008; Storlazzi et al., 2009). For these proximal stressors it is likely to be the duration and frequency caused by dredging


R. Jones et al / Marine Pollution Bulletin xxx (2015) xxx-xxx

that differs from natural conditions rather than the intensity as with sediment deposition. Corals are exposed to conditions (of low light and high SSCs) that they have experienced previously, in the short term, and are physiologically acclimated to these natural conditions. How they do this is via a range of strategies including photo-acclimatory changes, shifts from autotrophy to heterotrophy, and replenishment of energy reserves between turbidity events (Anthony, 2000, 2006; Anthony and Fabricius, 2000; Anthony and Hoegh-Guldberg, 2003; Anthony and Larcombe, 2000).

There are currently no suitable techniques for measuring low mg per cm2 deposition events with sufficient resolution to be effective as a monitoring tool for dredging programmes (Field et al., 2012; Perkey and Wadman, 2013; Risk and Edinger, 2011; Thomas and Ridd, 2004), although use ofupward pointing optical backscatter devices offer promise (Thomas and Ridd, 2005; Thomas et al., 2003). Once information becomes available on the range of sediment deposition rates (as mass per unit area or thickness deposition per day) over different time periods during natural conditions, or in the near- and far-field during dredging programmes, then this will allow contextualization of past studies and information that can be used for impact prediction purposes for sedimentation.

7.3. Seawater quality thresholds for coral reefs and future directions

Future ex situ studies need to clearly state what pressure parameter is being tested and recognise and eliminate the potential confounding effect of other parameters if values are to be proposed for threshold development. Studies should be conducted with locally collected reefal sediments rather than using sediment proxies such as carborundum. Studies to characterize the organic content of the sediments released into the water column by dredging (i.e. loss at the drag/cutter heads, by overflowing, or released by disposal at placement sites), and how it compares to surficial samples from the seabed before dredging are needed. Many studies have now shown the importance of the organic content of the sediments on coral sediment clearance and survivorship (Loiola et al., 2013; Weber et al., 2012; Weber et al., 2006). The use of low organic content or organic free sediments (Peters and Pilson, 1985; Riegl, 1995; Riegl and Branch, 1995), or proxies (Browne et al., 2014; Lui et al., 2012) has been suggested a starting point for understanding coral sediment shifting capabilities without additional variable of organic content, nutrient, and microbial content. Particle sizes should be measured and experiments examining the effects of sediment deposition in the near-field need to use up to coarse silt-sized sediments and finer fractions for far-field conditions. Contaminant levels need to be described if there is reasonable doubt they are of concern. PAR levels in tanks should be specified as maximum, as well as the daily light integral and recent advances in lighting technologies such as light emitting diodes (LEDs) means it is now possible to address spectral changes under plumes.

As Harris et al. (2014) warn in their review of principles of sound ecotoxicology, there is a danger associated with an incomplete understanding of exposure pathways and the use of conditions that are unrepresentative of the majority of situations typically encountered by wildlife. This statement seems applicable to studies on the effects of sediments on corals (see also Storlazzi et al., 2015; Storlazzi et al., 2011), as many experiments have been conducted without explicit justification of the exposure regimes especially with studies examining sedimentation. Harris et al. (2014) suggest authors should be open and honest about the context of their study to those conditions which have been measured (or predicted) in the real environment and the explanation of the exposure conditions for futures studies should be comprehensive. This will then enable judgements to be made of whether high suspended-sediment concentrations and the associated light reduction and sediment smothering at different distances from dredging is a hazard or a risk to underlying communities, and allow the development of

the water quality thresholds for impact prediction and monitoring purposes.

Author contributions

Conceived the study RJ, PB-B, RF. Conducted the analyses, and developed the model RJ, P B-B RJ. Conducted the PSD and light quality and quantity measurements MS, WK. All authors contributed to the writing and approved the final review.

Competing interests

The authors have declared no competing interests exist.


This project was funded by Chevron Australia, Woodside Energy Limited, BHP Billiton and Rio Tinto through contributions to the Western Australian Marine Science Institution (WAMSI) Dredging Science Node. This research was enabled by data and information generously provided by Woodside Energy Ltd, Rio Tinto Iron Ore and Chevron. The funders had no role in data analysis, decision to publish, or preparation of the manuscript.


Anthony, K., 1999. Coral suspension feeding on fine particulate matter. J. Exp. Mar. Biol. Ecol. 232, 85-106.

Anthony, K., 2000. Enhanced particle-feeding capacity of coral on turbid reefs (Great Barrier Reef, Australia). Coral Reefs 19,59-67. Anthony, K., 2006. Enhanced energy status of corals on coastal, high-turbidity reefs. Mar.

Ecol. Prog. Ser. 319,111-116. Anthony, K., Fabricius, K., 2000. Shifting roles of heterotrophy and autotrophy in coral energetics under varying turbidity. J. Exp. Mar. Biol. Ecol. 252, 221 -253. Anthony, K., Hoegh-Guldberg, O., 2003. Kinetics of photoacclimation in corals. Oecologia 134, 23-31.

Anthony, K., Larcombe, P., 2000. Coral reefs in turbid waters: sediment-induced stresses in corals and likely mechanisms of adaptation. Proceedings of the 9th International Coral Reef Symposium. Bali, Indonesia. 1, pp. 239-244. Anthony, K., Ridd, P., Orpin, A., Larcombe, P., Lough, J., 2004. Temporal variation of light availability in coastal benthic habitats: effects of clouds, turbidity, and tides. Limnol. Oceanogr. 49,2201-2211. Bailey-Serres, J., Voesenek, L.A.C.J., 2008. Flooding stress: acclimations and genetic diversity. Annu. Rev. Plant Biol. 59,313-339. Bainbridge, Z.T., Wolanski, E., Alvarez-Romero, J.G., Lewis, S.E., Brodie, J.E., 2012. Fine sediment and nutrient dynamics related to particle size and floc formation in a Burdekin River flood plume, Australia. Mar. Pollut. Bull. 65, 236-248. Bak, R.P.M., 1978. Lethal and sublethal effects of dredging on reef coral. Mar. Pollut. Bull. 9, 14-16.

Bak, R., Elgershuizen, J., 1976. Patterns of oil-sediment rejection in corals. Mar. Biol. 37, 105-113.

Barnes, D., 1973. Growth in colonial scleractinians. Bull. Mar. Sci. 23, 280-298. Bell, J., McGrath, E., Biggerstaff, A., Bates, T., Bennett, H., Marlow, J., Shaffer, M., 2015. Sediment impacts on marine sponges. Mar. Pollut. Bull. 94, 5-13. Bothner, M.H., Reynolds, R.L., Casso, MA, Storlazzi, C.D., Field, M.E., 2006. Quantity, composition, and source of sediment collected in sediment traps along the fringing coral reef off Molokai, Hawaii. Mar. Pollut. Bull. 52,1034-1047. Brown, B., LeTissier, M., Scoffin, T., Tudhope, A., 1990. Evaluation of the environmental impact of dredging on intertidal coral reefs at Ko Phuket, Thailand, using ecological and physiological parameters. Mar. Ecol. Prog. Ser. 65, 273-281. Browne, N.K., Precht, E., Last, K.S., Todd, PA., 2014. Photo-physiological costs associated with acute sediment stress events in three near-shore turbid water corals. Mar. Ecol. Prog. Ser. 502,129-143. Browne, N.K., Smithers, S.G., Perry, C.T., Ridd, P.V., 2012. A field-based technique for measuring sediment flux on coral reefs: application to turbid reefs on the Great Barrier Reef. J. Coast. Res. 284,1247-1262. Browne, N.K., Tay, J., Todd, P.A., 2015. Recreating pulsed turbidity events to determine coral-sediment thresholds for active management. J. Exp. Mar. Biol. Ecol. 466, 98-109.

Buesseler, K.O., Antia, A.N., Chen, M., Fowler, S.W., Gardner, W.D., Gustafsson, O., Harada, K., Michaels, A.F., der Loeff, M.R., Sarin, M., 2007. An assessment of the use of sediment traps for estimating upper ocean particle fluxes. J. Mar. Res. 65,345-416. Butman, C.A., Grant, W.D., Stolzenbach, K.D., 1986. Predictions of sediment trap biases in turbulent flows: a theoretical analysis based on observations from the literature. J. Mar. Res. 44,601-644. Chapman, P.M., 2002. Integrating toxicology and ecology: putting the "eco" into ecotoxi-cology. Mar. Pollut. Bull. 44, 7-15.


R.Jones et al./Marine Pollution Bulletin xxx (2015) xxx-xxx 19

Cooper, T.F., Fabricius, K.E., 2012. Pigmentation of massive corals as a simple bioindicator for marine water quality. Mar. Pollut Bull. 65,333-341.

Davies-Colley, R., Smith, D., 2001. Turbidity, suspended sediment and water quality: a review. J. Am. Water Works Assoc. 37,1085-1101.

Dennison, W.C., Orth, R.J., Moore, K.A., Stevenson, J.C., Carter, V., Kollar, S., Bergstrom, P.W., Batiuk, RA, 1993. Assessing water quality with submersed aquatic vegetation. Bioscience 43, 86-94.

DeSalvo, M., Estrada, A., Sunagawa, S., Medina, M., 2012. Transcriptomic responses to darkness stress point to common coral bleaching mechanisms. Coral Reefs 31, 215-228.

DEWHA, 2009. National Assessment Guidelines Dredging (2009). In: W. Heritage, the Arts (DEWHA) (Eds.), Department of the Environment (http://www.environment, Canberra, Australia).

Dodge, R.E., Vaisnys, J.R., 1977. Coral populations and growth patterns: responses to sedimentation and turbidity associated with dredging. J. Mar. Res. 35, 715-730.

Dubinsky, Z., Falkowski, P., Porter, J., Muscatine, L., 1984. Absorption and utilization of radiant energy by light-and shade-adapted colonies of the hermatypic coral Stylophora pistillata. Proc. R Soc. Lond. Ser. B Biol. Sci. 222, 203-214.

Duclos, P.-A., Lafite, R., Le Bot, S., Rivoalen, E., Cuvilliez, A., 2013. Dynamics of turbid plumes generated by marine aggregate dredging: an example of a macrotidal environment (the Bay of Seine, France). J. Coast. Res. 25-37.

Eggleton, J., Thomas, K.V., 2004. A review of factors affecting the release and bioavailability of contaminants during sediment disturbance events. Environ. Int. 30, 973-980.

Ellington, R., 1977. Aerobic and anaerobic degradation of glucose by the estuarine sea anemone, Diadumene leucolena. Comp. Biochem. Physiol. B: Biochem. Mol. Biol. 58, 173-175.

Ellington, W., 1980. Some aspects of the metabolism of the sea-anemone Haliplanella luciae (Verrill) during air exposure and hypoxia. 1,255-262.

Ellington, W., 1982. Metabolic responses of the sea anemone Bunodosoma cavernata (Bosc) to declining oxygen tensions and anoxia. Physiol. Zool. 55, 240-249.

Enriquez, S., Méndez, E.R., Iglesias-Prieto, R., 2005. Multiple scattering on coral skeletons enhances light absorption by symbiotic algae. Limnol. Oceanogr. 50,1025-1032.

EPA, 2011. Environmental Assessment Guidleine for Marine Dredging Programs EAG7. Environmental Protection Authority (EPA), Perth, Western Australia, p. 36.

Erftemeijer, P.L., Riegl, B., Hoeksema, B.W., Todd, P.A., 2012. Environmental impacts of dredging and other sediment disturbances on corals: a review. Mar. Pollut. Bull. 64, 1737-1765.

Fabricius, K.E., 2005. Effects of terrestrial runoff on the ecology of corals and coral reefs: review and synthesis. Mar. Pollut. Bull. 50,125-146.

Fabricius, K.E., Wolanski, E., 2000. Rapid smothering of coral reef organisms by muddy marine snow. Estuar. Coast. Shelf Sci. 50,115-120.

Falkenberg, L.J., Styan, C.A., 2014. Too much data is never enough: a review of the mismatch between scales of water quality data collection and reporting from recent marine dredging programmes. Ecol. Indic. 45, 529-537.

Falkowski, P.G., Dubinsky, Z., 1981. Light-shade adaptations of Stylophora pistillata, a hermatypic corals from the Gulf of Eilat. Nature 289,172-174.

Falkowski, P.G., Dubinsky, Z., Muscatine, L., Porter, J.W., 1984. Light and the bioenergetics of a symbiotic coral. Bioscience 34, 705-709.

Ferrier-Pages, C., Witting, J., Tambutté, E., Sebens, K., 2003. Effect of natural zooplankton feeding on the tissue and skeletal growth of the scleractinian coral Stylophora pistillata. Coral Reefs 22, 229-240.

Field, M.E., Chezar, H., Storlazzi, C.D., 2012. SedPods: a low-cost coral proxy for measuring net sedimentation. Coral Reefs 32,155-159.

Fisher, R., Stark, C., Ridd, P., Jones, R., 2015. Spatial patterns in water quality changes during dredging in tropical environments. PLoS ONE (in press).

Flores, F., Hoogenboom, M.O., Smith, L.D., Cooper, T.F., Abrego, D., Negri, A.P., 2012. Chronic exposure of corals to fine sediments: lethal and sub-lethal impacts. PLoS One 7, e37795.

Foster, T., Corcoran, E., Erftemeijer, P., Fletcher, C., Peirs, K., Dolmans, C., Smith, A., Yamamoto, H., Jury, M., 2010. Dredging and port construction around coral reefs. PIANC Environmental Commission, Report No 108.

Freudenthal, H., 1962. Symbiodinium gen. nov. and Symbiodinium microadriaticum sp. nov., a Zooxanthella: taxonomy, life cycle, and morphology. J. Protozool. 9,45-52.

Fukao, T., Bailey-Serres, J., 2004. Plant responses to hypoxia — is survival a balancing act? Trends Plant Sci. 9,449-456.

Gardner, W., 1980. Field assessment of sediment traps. J. Mar. Res. 38,41-52.

Gardner, W., Richardson, M., Hinga, K., Biscaye, P., 1983. Resuspension measured with sediment traps in a high-energy environment. Earth Planet. Sci. Lett. 66, 262-278.

Gattuso, J.-P., Allemand, D., Frankignoulle, M., 1999. Photosynthesis and calcification at cellular, organismal and community levels in coral reefs: a review on interactions and control by carbonate chemistry. Am. Zool. 39,160-183.

Gleason, D.F., 1998. Sedimentation and distributions of green and brown morphs of the Caribbean coral Porites astreoides Lamarck. J. Exp. Mar. Biol. Ecol. 230, 73-89.

Goreau, T.F., 1959. The physiology of skeleton formation in corals. I. A method for measuring the rate of calcium deposition by corals under different conditions. Biol. Bull. Mar. Biol. Lab. Woods Hole 59-75.

Goreau, T.F., Goreau, N.I., Yonge, C., 1971. Reef corals: autotrophs or heterotrophs? Biol. Bull. Mar. Biol. Lab. Woods Hole 141,247-260.

Halldal, P., 1968. Photosynthetic capacities and photosynthetic action spectra of endozoic algae of the massive coral Favia. Biol. Bull. 134,411-424.

Harris, CA, Scott, A.P., Johnson, A.C., Panter, G.H., Sheahan, D., Roberts, M., Sumpter, J.P., 2014. Principles of sound ecotoxicology. Environ. Sci. Technol. 48, 3100-3111.

Hedge, L.H., Knott, N.A., Johnston, E.L., 2009. Dredging related metal bioaccumulation in oysters. Mar. Pollut. Bull. 58,832-840.

Hodgson, G., 1990. Tetracycline reduces sedimentation damage to corals. Mar. Biol. 104, 493-496.

Houlbreque, F., Ferrier-Pages, C., 2009. Heterotrophy in tropical scleractinian corals. Biol. Rev. Camb. Philos. Soc. 84,1-17.

Hubbard, JA, Pocock, Y.P., 1972. Sediment rejection by recent scleractinian corals: a key to palaeo-environmental reconstruction. Geol. Rundsch. 61,598-626.

Jerlov, N.G., 1976. Marine Optics. Elsevier.

Jing, L., Ridd, P.V., 1996. Wave-current bottom shear stresses and sediment resuspension in Cleveland Bay, Australia. Coast. Eng. 29,169-186.

Jones, R.J., 2008. Coral bleaching, bleaching-induced mortality, and the adaptive significance of the bleaching response. Mar. Biol. 154, 65-80.

Jones, R.J., 2011a. Spatial patterns of chemical contamination (metals, PAHs, PCBs, PCDDs/ PCDFS) in sediments of a non-industrialized but densely populated coral atoll/small island state (Bermuda). Mar. Pollut. Bull.

Jones, R., 2011b. Environmental effects of the cruise tourism boom: sediment resuspension from cruise ships and the possible effects of increased turbidity and sediment deposition on corals (Bermuda). Bull. Mar. Sci. 87, 659-679.

Jones, R., Hoegh-Guldberg, O., 2001. Diurnal changes in the photochemical efficiency of the symbiotic dinoflagellates (Dinophyceae) of corals: photoprotection, photoinactivation and the relationship to coral bleaching. Plant Cell Environ. 24, 89-99.

Jones, R., Fisher, R., Stark, C., Ridd, P., 2015a. Temporal patterns in water quality from dredging in tropical environments. PLoS One 10 (10), e0137112. 10.1371/journal.pone.0137112.

Jones, R., Ricardo, G., AP, N., 2015b. Effects of sediments on the reproductive cycle of corals. Mar. Pollut. Bull. xxx-xxx

Junjie, R.K., Browne, N.K., Erftemeijer, P.L.A., Todd, P.A., 2014. Impacts of sediments on coral energetics: partitioning the effects of turbidity and settling particles. PLoS One 9, e107195.

Jurg, B., 1996. Towards a new generation of sediment traps and a better measurement/ understanding of settling particle flux in lakes and oceans: a hydrodynamical protocol. Aquat. Sci. 58, 283-296.

Kendall, J., Powell, E., Connor, S., Bright, T., 1983. The effects of drilling fluids (muds) and turbidity on the growth and metabolic state of the coral Acropora cervicornis, with comments on methods of normalization for coral data. Bull. Mar. Sci. 33, 336-352.

Kirk, J.T., 1985. Effects of Suspensoids (Turbidity) on Penetration of Solar Radiation in Aquatic Ecosystems, Perspectives in Southern Hemisphere Limnology. Springer, 208, p. 195.

Kirk, J., 1994. Light and Photosynthesis in Aquatic Ecosystems. Third Edition. Third edition. Cambridge University Press, New York.

Koskela, R.W., Ringeltaube, P., Small, A.R., Koskela, T.V., Fraser, A.R., Lee, J.D., Marshall, P., 2002. Using predictive monitoring to mitigate construction impacts in sensitive marine environments. 10th Pacific Congress on Marine Science and Technology (PACON 2002) July 21-26, 2002, Chiba, Japan.

Kozerski, H.-P., 1994. Possibilities and limitations of sediment traps to measure sedimentation and resuspension. Hydrobiologia 284, 93-100.

Kuhl, M., Cohen, Y., Dalsgaard, T., Jorgensen, B.B., Revsbech, N.P., 1995. Microenvironment and photosynthesis of zooxanthellae in scleractinian corals studied with microsensors for O2, pH and light. Mar. Ecol. Prog. Ser. 117,159-172.

LaBarbera, M., 1984. Feeding currents and particle capture mechanisms in suspension feeding animals. Am. Zool. 24, 71-84.

Larcombe, P., Woolfe, K., 1999. Increased sediment supply to the Great Barrier Reef will not increase sediment accumulation at most coral reefs. Coral Reefs 18, 163-169.

Larcombe, P., Costen, A., Woolfe, K.J., 2001. The hydrodynamic and sedimentary setting of nearshore coral reefs, central Great Barrier Reef shelf, Australia: Paluma Shoals, a case study. Sedimentology 48, 811-835.

Larcombe, P., Ridd, P., Prytz, A., Wilson, B., 1995. Factors controlling suspended sediment on inner-shelf coral reefs, Townsville, Australia. Coral Reefs 14,163-171.

Lasker, H.R., 1980. Sediment rejection by reef corals: the roles of behavior and morphology in Montastrea cavernosa (Linnaeus). J. Exp. Mar. Biol. Ecol. 47, 77-87.

Lawrence, D., Dagg, M.J., Liu, H., Cummings, S.R., Ortner, P.B., Kelble, C., 2004. Wind events and benthic-pelagic coupling in a shallow subtropical bay in Florida. Mar. Ecol. Prog. Ser. 266,1-13.

Lesser, M.P., 2004. Experimental biology of coral reef ecosystems. J. Exp. Mar. Biol. Ecol. 300, 217-252.

Levy, O., Dubinsky, Z., Achituv, Y., 2003. Photobehavior of stony corals: responses to light spectra and intensity. J. Exp. Biol. 206,4041-4049.

Lewis, J.B., 1976. Experimental tests of suspension feeding in Atlantic reef corals. Mar. Biol. 36,147-150.

Lewis, J., Price, W., 1975. Feeding mechanisms and feeding strategies of Atlantic reef corals. J. Zool. 176, 527-544.

Lewis, J., Price, W., 1976. Patterns of ciliary currents in Atlantic reef corals and their functional significance. J. Zool. 178, 77-89.

Lirman, D., Herlan, J., Langdon, C., Capo, T., 2008. To nutrient-enriched sediments mitigates the negative impacts of sedimentation on coral growth in the Caribbean corals Porites astreoides and Siderastrea siderea. Proceedings of the 11th International Coral Reef Symposium. Fort Lauderdale, Florida. US, pp. 900-904.

Logan, A., 1988. Sediment-shifting capability in the recent solitary coral Scolymia cubensis (Milne-Edwards and Haime) from Bermuda. Bull. Mar. Sci. 43,241-248.

Logan, A., Yang, L., Tomascik, T., 1994. Linear skeletal extension rates in two species of Diploria from high-latitude reefs in Bermuda. Coral Reefs 13, 225-230.

Loiola, M., Oliveira, M.D.M., Kikuchi, R.K.P., 2013. Tolerance of Brazilian brain coral Mussismilia braziliensis to sediment and organic matter inputs. Mar. Pollut. Bull. 77, 55-62.

Lopez, G.R., Levinton, J.S., 1987. Ecology of deposit-feeding animals in marine sediments. Q.Rev. Biol. 235-260.


20 R. Jones et al. / Marine Pollution Bulletin xxx (2015) xxx-xxx

Loya, Y., 1976. Effects of water turbidity and sedimentation on the community structure of Puerto Rican corals. Bull. Mar. Sci. 26,450-466.

Lui, G.C., Setiawan, W., Todd, P., Erftemeijer, P.L., 2012. Among genotype variation for sediment rejection in the reef building coral Diploastrea heliopora (Lamark, 1816). Raffles Bull. Zool. 60, 525-531.

Madden, RA, Julian, P.R., 1994. Observations of the 40-50-day tropical oscillation — a review. Mon. Weather Rev. 122, 814-837.

Mapstone, B.D., Choat, J., Cumming, R., Oxley, W., 1992. The fringing reefs of Magnetic Island: benthic biota and sedimentation-a baseline study: a report to the Great Barrier Reef Marine Park Authority. Research report No.13. Great Barrier Reef Marine Park Authority, Townsville, Australia.

Marcelino, L.A., Westneat, M.W., Stoyneva, V., Henss, J., Rogers, J.D., Radosevich, A., Turzhitsky, V., Siple, M., Fang, A., Swain, T.D., Fung, J., Backman, V., 2013. Modulation of light-enhancement to symbiotic algae by light-scattering in corals and evolutionary trends in bleaching. PLoS One 8, e61492.

Marshall, N., 1965. Detritus over the reef and its potential contribution to adjacent waters of Eniwetok Atoll. Ecology 46, 343-344.

Marshall, S.M., Orr, A.P., 1931. Sedimentation on Low Isles Reef and its relation to coral growth. Brit. Mus. (Nat. Hist.).

Mass, T., Kline, D.I., Roopin, M., Veal, C.J., Cohen, S., Iluz, D., Levy, O., 2010. The spectral quality of light is a key driver of photosynthesis and photoadaptation in Stylophora pistillata colonies from different depths in the Red Sea. J. Exp. Biol. 213,4084-4091.

Masselink, G., Hughes, M., Knight, J., 2014. Introduction to Coastal Processes and Geomor-phology. Routledge.

McAnally, W.H., Mehta, A.J., 2001. Coastal and Estuarine Fine Sediment Processes. Elsevier Science.

McClanahan, T., Obura, D., 1997. Sedimentation effects on shallow coral communities in Kenya. J. Exp. Mar. Biol. Ecol. 209,103-122.

Meyer, J.L., Schultz, E.T., 1985. Tissue condition and growth rate of corals associated with schooling fish. Limnol. Oceanogr. 30,157-166.

Mills, M., Sebens, K., 1997. Particle ingestion efficiency of the corals Siderastrea siderea and Agaricia agaricites: effects of flow speed and sediment loads. Proceedings of the 8th International Coral Reef Symposium. Panama 2,1059-1064.

Mills, M.M., Sebens, K.P., 2004. Ingestion and assimilation of nitrogen from benthic sediments by three species of coral. Mar. Biol. 145,1097-1106.

Mills, M.M., Lipschultz, F., Sebens, K.P., 2004. Particulate matter ingestion and associated nitrogen uptake by four species of scleractinian corals. Coral Reefs 23, 311-323.

Morel, A., 1978. Available, usable, and stored radiant energy in relation to marine photosynthesis. Deep-Sea Res. 25, 673-688.

Morgan, B., Rate, A.W., Burton, E.D., 2012. Water chemistry and nutrient release during the resuspension of FeS-rich sediments in a eutrophic estuarine system. Sci. Total Environ. 432, 47-56.

Muscatine, L., 1990. The role of symbiotic algae in carbon and energy flux in reef corals. Ecosyst. World 25, 75-87.

Muscatine, L., Porter, J.W., 1977. Reef corals: mutualistic symbioses adapted to nutrient-poor environments. Bioscience 27,454-460.

Muscatine, L., Ferrier-Pagés, C., Blackburn, A., Gates, R.D., Baghdasarian, G., Allemand, D., 1998. Cell-specific density of symbiotic dinoflagellates in tropical anthozoans. Coral Reefs 17, 329-337.

Norton, S.B., Cormier, S.M., Suter II, G.W., Schofield, K., Yuan, L., Shaw-Allen, P., Ziegler, C.R., 2009. CADDIS: The Causal Analysis/diagnosis Decision Information System, Decision Support Systems for Risk-based Management of Contaminated Sites. Springer, pp. 1-24.

Ogston, A.S., Field, M.E., 2010. Predictions of turbidity due to enhanced sediment resuspension resulting from sea-level rise on a fringing coral reef: evidence from Molokai, Hawaii. J. Coast. Res. 26,1027-1037.

Ogston, A.S., Storlazzi, C.D., Field, M.E., Presto, M.K., 2004. Sediment resuspension and transport patterns on a fringing reef flat, Molokai, Hawaii. Coral Reefs 23, 559-569.

Orpin, A.R., Ridd, P.V., 2012. Exposure of inshore corals to suspended sediments due to wave-resuspension and river plumes in the central Great Barrier Reef: a reappraisal. Cont. Shelf Res. 47,55-67.

Orpin, A., Ridd, P., Thomas, S., Anthony, K., Marshall, P., Oliver, J., 2004. Natural turbidity variability and weather forecasts in risk management of anthropogenic sediment discharge near sensitive environments. Mar. Pollut. Bull. 49, 602-612.

Perkey, D.W., Wadman, H.M., 2013. Laboratory Feasibility Study Concerning the Use of the SediMeter™ to Detect Fine-scale (<1 mm) Sedimentation Resulting From Dredging Operations. DOER Technical Notes Collection. ERDC TN-DOER-T11. US Army Engineer Research and Development Center, Vicksburg, MS ( mil/dots/doer/).

Perry, C., Taylor, K., 2009. Environmental Sedimentology. John Wiley & Sons.

Peters, E.C., Pilson, M.E., 1985. A comparative study of the effects of sedimentation on symbiotic and asymbiotic colonies of the coral Astrangia danae Milne Edwards and Haime 1849. J. Exp. Mar. Biol. Ecol. 92,215-230.

Philipp, E., Fabricius, K., 2003. Photophysiological stress in scleractinian corals in response to short-term sedimentation. J. Exp. Mar. Biol. Ecol. 287, 57-78.

Piniak G.A., 2007. Effects of two sediment types on the fluorescence yield of two Hawaiian scleractinian corals. Mar. Environ. Res. 64, 456-468.

R Core Team, 2014. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria (URL

Reef, R., Kaniewska, P., Hoegh-Guldberg, O., 2009. Coral skeletons defend against ultraviolet radiation. PLoS One 4, e7995.

Reynolds, C.S., Wiseman, S.W., Gardner, W., 1980. An annotated bibliography of aquatic sediment traps and trapping methods. Freshwater Biological Association. Occasional Publication No.11 (54 pp.).

Rice, S., Hunter, C., 1992. Effects of suspended sediment and burial on scleractinian corals from west central Florida patch reefs. Bull. Mar. Sci. 51,429-442.

Riegl, B., 1995. Effects of sand deposition on scleractinian and alcyonacean corals. Mar. Biol. 121,517-526.

Riegl, B., Branch, G., 1995. Effects of sediment on the energy budgets of four scleractinian (Bourne 1900) and five alcyonacean (Lamouroux 1816) corals. J. Exp. Mar. Biol. Ecol. 186, 259-275.

van Rijn, L.C., 2007. Unified view of sediment transport by currents and waves. I: initiation of motion, bed roughness, and bed-load transport. J. Hydraul. Eng. 133,649-667.

Risk M.J., Edinger, E., 2011. Impacts of sediment on coral reefs. In: Hopley, D. (Ed.), Encyclopedia of Modern Coral Reefs. Springer, Netherlands, pp. 575-586.

Rogers, C.S., 1979. The effect of shading on coral reef structure and function. J. Exp. Mar. Biol. Ecol. 41, 269-288.

Rogers, C.S., 1990. Responses of coral reefs and reef organisms to sedimentation. Mar. Ecol. Prog. Ser. 62,185-202.

Rosenfeld, M., Bresler, V., Abelson, A., 1999. Sediment as a possible source of food for corals. Ecol. Lett. 2, 345-348.

Roth, M.S., 2014. The engine of the reef: photobiology of the coral-algal symbiosis. Front. Microbiol. 5, 1 -22.

Roth, E., Jeon, K., Stacey, G., 1988. Homology in endosymbiotic systems: the term 'symbiosome'. In: Palacios, R., Verma, D.P.S. (Eds.), Molecular Genetics of Plant-Microbe Interactions. St. Paul, MN, pp. 220-225.

Rubenstein, D.I., Koehl, M., 1977. The mechanisms of filter feeding: some theoretical considerations. Am. Nat. 111, 981-994.

Schuhmacher, H., 1974. On the conditions accompanying the first settlement of corals on artificial reefs with special reference to the influence of grazing sea urchins (Eilat, Red Sea). Proceedings of the 2nd International Coral Reef Symposium, Brisbane, Australia. 1, pp. 257-267.

Schuhmacher, H., 1977. Ability of fungiid corals to overcome sedimentation. Proceedings of the 3rd International Coral Reef Symposium, Miami, US, pp. 503-510.

Sebens, K., Grace, S., Helmuth, B., Maney Jr., E., Miles, J., 1998. Water flow and prey capture by three scleractinian corals, Madracis mirabilis, Montastrea cavernosa and Porites porites, in a field enclosure. Mar. Biol. 131, 347-360.

Sebens, K., Vandersall, K., Savina, L., Graham, K., 1996. Zooplankton capture by two scleractinian corals, Madracis mirabilis and Montastrea cavernosa, in a field enclosure. Mar. Biol. 127, 303-317.

Shapiro, O.H., Fernandez, V.I., Garren, M., Guasto, J.S., Debaillon-Vesque, F.P., Kramarsky-Winter, E., Vardi, A., Stocker, R., 2014. Vortical ciliary flows actively enhance mass transport in reef corals. Proc. Natl. Acad. Sci. 111,13391-13396.

Shashar, N., Kinane, S., Jokiel, P.L., Patterson, M.R., 1996. Hydromechanical boundary layers over a coral reef. J. Exp. Mar. Biol. Ecol. 199,17-28.

Shick, J.M., 1991. Energy metabolism and respiratory gas exchange, a functional biology of sea anemones. Springer, pp. 119-173.

Shimeta, J., Koehl, M.A.R., 1997. Mechanisms of particle selection by tentaculate suspension feeders during encounter, retention, and handling. J. Exp. Mar. Biol. Ecol. 209, 47-73.

Sofonia, J., Anthony, K., 2008. High-sediment tolerance in the reef coral Turbinaria mesenterina from the inner Great Barrier Reef lagoon (Australia). Estuar. Coast. Shelf Sci. 78, 748-752.

Spearman, J., 2015. A review of the physical impacts of sediment dispersion from aggregate dredging. Mar. Pollut. Bull. 94, 260-277.

Spearman, J., Bray, R., Land, J., Burt, T., Mead, C., Scott, D., 2007. Plume dispersion modelling using dynamic representation of trailer dredger source terms. Proc. Mar. Sci. 8, 417-448.

Stafford-Smith, M.G., 1993. Sediment-rejection efficiency of 22 species of Australian scleractinian corals. Mar. Biol. 115, 229-243.

Stafford-Smith, M., Ormond, R., 1992. Sediment-rejection mechanisms of 42 species of Australian scleractinian corals. Mar. Freshw. Res. 43, 683-705.

Stat, M., Baker, A.C., Bourne, D.G., Correa, A.M.S., Forsman, Z., Huggett, M.J., Pochon, X., Skillings, D., Toonen, RJ., van Oppen, M.J.H., Gates, R.D., 2012. Chapter one — molecular delineation of species in the coral holobiont. In: Michael, L. (Ed.)Adv. Mar. Biol.Academic Press, pp. 1 -65

Storlazzi, C.D., Jaffe, B.E., 2008. The relative contribution of processes driving variability in flow, shear, and turbidity over a fringing coral reef: West Maui, Hawaii. Estuar. Coast. Shelf Sci. 77, 549-564.

Storlazzi, C.D., Field, M.E., Bothner, M.H., 2011. The use (and misuse) of sediment traps in coral reef environments: theory, observations, and suggested protocols. Coral Reefs 30, 23-38.

Storlazzi, C.D., Field, M.E., Bothner, M.H., Presto, M.K., Draut, A.E., 2009. Sedimentation processes in a coral reef embayment: Hanalei Bay, Kauai. Mar. Geol. 264,140-151.

Storlazzi, C., Norris, B., Rosenberger, K., 2015. The influence of grain size, grain color, and suspended-sediment concentration on light attenuation: why fine-grained terrestrial sediment is bad for coral reef ecosystems. Coral Reefs 1-9.

Storlazzi, C.D., Ogston, A.S., Bothner, M.H., Field, M.E., Presto, M.K., 2004. Wave- and tidally-driven flow and sediment flux across a fringing coral reef: Southern Molokai, Hawaii. Cont. Shelf Res. 24,1397-1419.

Su, S., Pearlman, L., Rothrock, J., Iannuzzi, T., Finley, B., 2002. Potential long-term ecological impacts caused by disturbance of contaminated sediments: a case study. Environ. Manag. 29, 234-249.

Szabo, M., Wangpraseurt, D., Tamburic, B., Larkum, A.W.D., Schreiber, U., Suggett, D.J., Kühl, M., Ralph, P.J., 2014. Effective light absorption and absolute electron transport rates in the coral Pocillopora damicornis. Plant Physiol. Biochem. 83,159-167.

Szmant-Froelich, A., Johnson, V., Hoehn, T., Battey, J., Smith, G., Fleischmann, E., Porter, J., Dallmeyer, D., 1981. The physiological effects of oil drilling muds on the Caribbean coral Montastrea annularis. Proceedings of the 4th International Coral Reef Symposium, Manila, Philippines, pp. 163-168.

Telesnicki, G.J., Goldberg, W.M., 1995. Effects of turbidity on the photosynthesis and respiration of two South Florida reef coral species. Bull. Mar. Sci. 57, 527-539.


R.Jones et al./Marine Pollution Bulletin xxx (2015) xxx-xxx 21

Thomas, S., Ridd, P.V., 2004. Review of methods to measure short time scale sediment accumulation. Mar. Geol. 207,95-114.

Thomas, S., Ridd, P., 2005. Field assessment of innovative sensor for monitoring of sediment accumulation at inshore coral reefs. Mar. Pollut. Bull. 51, 470-480.

Thomas, S., Ridd, P.V., Renagi, O., 2003. Laboratory investigation on the effect of particle size, water flow and bottom surface roughness upon the response of an upward-pointing optical backscatter sensor to sediment accumulation. Cont. Shelf Res. 23, 1545-1557.

Thompson, J.H., 1980. Responses of Selected Scleractinian Corals to Drilling Fluids Used in the Marine Environment (PhD Dissertation) Texas A & M University, College Station.

Thompson, J.H., Shinn, E.A., Bright, T.J., 1980. Effects of drilling mud on seven species of reef-building corals as measured in the field and laboratory. Elsevier Oceanogr. Ser. 27,433-453.

Todd, P.A., Sidle, R.C., Lewin-Koh, N.J.I., 2004. An aquarium experiment for identifying the physical factors inducing morphological change in two massive scleractinian corals. J. Exp. Mar. Biol. Ecol. 299, 97-113.

Tomascik, T., Sander, F., 1985. Effects of eutrophication on reef-building corals. I. Growth rate of the reef-building coral Montastrea annularis. Mar. Biol. 87,143-155.

Tuszynski,J., 2013. caTools: Tools: moving window statistics, GIF, Base64, ROC AUC, etc. R package version 1.16. (

Tyler, J., 1966. Report on the second meeting of the joint group of experts on photosyn-thetic radiant energy. UNESCO Tech. Pap. Mar. Sci. 2,1-11.

USEPA, 2004. In: Assessment, N.C.f.E. (Ed.), Development plan for the causal/diagnosis decision information system. USEPA, Washington, DC.

Van Lancker, V., Baeye, M., 2015. Wave glider monitoring of sediment transport and dredge plumes in a shallow marine sandbank environment. PLoS One 10, e0128948.

Van Rijn, L.C., van Rijn, L.C., van Rijn, L.C., 1990. Principles of Fluid Flow and Surface Waves in Rivers, Estuaries, Seas and Oceans. Aqua Publications Amsterdam, The Netherlands.

Vaughan, T.W., 1916. The results of investigations of the ecology of the Floridian and Bahaman Shoal-Water Corals. Proc. Natl. Acad. Sci. U. S. A. 2, 95-100.

VBKO, 2003. Protocol for the field measurementof sediment release from dredgers. A practical guide to measuring sediment release from dredging plant for calibration and verification of numerical models. HR Wallingford Ltd & Dredging Research Ltd. Reporrt produced for VBKO TASS project by. (83 pp.).

Verspecht, F., Pattiaratchi, C., 2010. On the significance of wind event frequency for partic-ulate resuspension and light attenuation in coastal waters. Cont. Shelf Res. 30, 1971-1982.

Wakefield, T., Kempf, S., 2001. Development of host- and symbiont-specific monoclonal antibodies and confirmation of the origin of the symbiosome membrane in a Cnidar-ian-diniflagellate symbiosis. Biol. Bull. Mar. Biol. Lab. Woods Hole 200,127-143.

Wangpraseurt, D., Larkum, A.W., Ralph, P.J., Kühl, M., 2012. Light gradients and optical microniches in coral tissues. Front. Microbiol. 3.

Wangpraseurt, D., Tamburic, B., Szabo, M., Suggett, D., Ralph, P.J., Kühl, M., 2014. Spectral effects on Symbiodinium photobiology studied with a programmable light engine. PLoS One 9, e112809.

Weber, M., de Beer, D., Lott, C., Polerecky, L., Kohls, K., Abed, R.M., Ferdelman, T.G., Fabricius, K.E., 2012. Mechanisms of damage to corals exposed to sedimentation. Proc. Natl. Acad. Sci. U. S. A. 109, E1558-E1567.

Weber, M., Lott, C., Fabricius, K.E., 2006. Sedimentation stress in a scleractinian coral exposed to terrestrial and marine sediments with contrasting physical, organic and geo-chemical properties. J. Exp. Mar. Biol. Ecol. 336, 18-32.

Wijgerde, T., Henkemans, P., Osinga, R., 2012. Effects of irradiance and light spectrum on growth of the scleractinian coral Galaxea fascicularis — applicability of LEP and LED lighting to coral aquaculture. Aquaculture 344,188-193.

Wild, C., Huettel, M., Klueter, A., Kremb, S.G., Rasheed, M.Y., Jorgensen, B.B., 2004. Coral mucus functions as an energy carrier and particle trap in the reef ecosystem. Nature 428, 66-70.

Wooldridge, S., 2013. A new conceptual model of coral biomineralisation: hypoxia as the physiological driver of skeletal extension. Biogeosciences 10, 2867-2884.

Wright, W., 1997. Tropical-extratropical cloudbands and Australian rainfall: I. Climatol. Int. J. Climatol. 17, 807-829.

Yeh, N., Yeh, P., Shih, N., Byadgi, O., Chih Cheng, T., 2014. Applications of light-emitting diodes in researches conducted in aquatic environment. Renew. Sust. Energ. Rev. 32, 611-618.

Yonge, C., 1930. Great Barrier Reef Expedition 1928-29, Scientific Reports. British Museum (Natural History), London (UK), pp. 13-57.

Yonge, C.M., Nicholls, A., 1931. Studies on the physiology of corals. V. The Effects of Starvation in Light and in Darkness on the Relationship Between Corals and ZooxanthellaeGreat Barrier Reef Expedition 1928-29, Scientific Reports. British Museum (Natural History), London (UK), pp. 13-57 (British Museum 1).

Zeileis, A., Grothendieck, G., 2005. Zoo: S3 infrastructure for regular and irregular time series. J. Stat. Softw. 14,1-27.