Mar Biol (2011) 158:1301–1310
DOI 10.1007/s00227-011-1649-z
ORIGINAL PAPER
Effects of sedimentation on the gorgonian Subergorgia suberosa
(Pallas, 1766)
Li-Chun Tseng • Hans-Uwe Dahms •
Nan-Jung Hsu • Jiang-Shiou Hwang
Received: 30 July 2009 / Accepted: 16 February 2011 / Published online: 8 March 2011
Ó Springer-Verlag 2011
Abstract Effects of oceanic and terrestrial sediment sus-
pensions on the behavioral traits activity, colony expansion,
and mucus formation of the gorgonian octocoral Suber-
gorgia suberosa (Pallas, 1766) were studied in the labora-
tory. Increased sediment concentration significantly
changed the logits of activity indices (as colony expansion)
and was correlated with decreased activity. For both sources
of sediments, the activity index decreased as the concen-
tration of sediment increased. The level of decrease was
more significance for terrestrial sediments. The activity
index showed no clear time pattern for the control group,
but a decrease over time occurred in both sediment treat-
ment groups. Activity change was linearly correlated with
exposure time and concentration of terrestrial and ocean
sediments. The activity indices of polyps were negatively
correlated with the formation of mucus sheets (correla-
tion = -0.536, p = 0.048). For both sources of sediments,
the frequency of mucus sheet formation (MSF) increased
with sediment concentration. Increases in terrestrial
Communicated by U. Sommer.
L.-C. Tseng  J.-S. Hwang (&)
Institute of Marine Biology, College of Life Sciences,
National Taiwan Ocean University, 2-Pei-Ning Road,
Chi-lung 202, Taiwan, ROC
e-mail: jshwang@mail.ntou.edu.tw
L.-C. Tseng
e-mail: lichun@mail.bestredeye.org
H.-U. Dahms
Green Life Science Department, Sangmyung University,
7 Hongji-dong, Jongno-gu, Seoul 110-743, Korea
N.-J. Hsu
Institute of Statistics, National Tsing-Hua University,
101 Kuang-Fu Road, Section 2, Hsin-Chu 30043, Taiwan, ROC
sediment loads resulted in more striking effects. The
probability for MSF under different sediment groups at
different concentrations over time was modeled by regres-
sions. Frequency of polyp activity is suggested as a suitable
biomarker to monitor the effects of sedimentation stress.
Introduction
Reefs around the tropical belt are globally affected by a
loss of biodiversity and vitality due to several stressors in
coastal environments (Wilkinson 2004; Anthony et al.
2007). Algal overgrowth (Hughes et al. 2003) associated
with overfishing (Pandolfi et al. 2003) and bleaching
associated with global warming (Hughes et al. 2003) are
widely accepted as major contributors to the catastrophic
decline of coral abundance and diversity. Coral diseases are
suggested as an additional reason for bleaching (Richard-
son 1998; Anthony et al. 2007), secondary to damage of
coral immune response (Sutherland et al. 2004). The role of
dissolved organic carbon (DOC—Kline et al. 2006),
nutrients (Ferrier-Pages et al. 2000), pollutants (Harrington
et al. 2005), and the role of sedimentation (Fabricius 2005;
Stamski and Field 2006; Wolanski et al. 2008; Takesue
et al. 2009) needs to be further investigated. Increasing
terrestrial runoff of sediments, nutrients and chemical
pollutants into coastal ecosystems causes degradation of
coastal coral reefs (Brodie et al. 2001; Presto et al. 2006).
Particularly, the harmful effects of sudden large or small
amounts of long-term deposits of terrigenous material on
coastal benthic communities (Lohrer et al. 2006) and coral
reefs (Fabricius et al. 2003a, b; Wilkinson 2004; Victor
et al. 2006; Wolanski et al. 2009).
For the purposes of this study, we defined sediment as
any particulate matter suspended and eventually settling in
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1302
water, including inorganic and organic matter or a com-
bination of both. Although the effects of turbidity and
sedimentation on coral reefs are of major concern world-
wide, little is known concerning the sources of sediments
affecting corals, and the effects of the amount and duration
of suspended sediment. We investigated the effects of
terrestrial and oceanic sediments on the behavior of the
gorgonian soft coral Subergorgia suberosa (Pallas, 1766)
using different concentrations of a defined grain size
fraction. Several behavioral aspects were tested for their
usefulness as indicators, including patterns of activity,
mucus production, and expansion behavior.
Materials and methods
Coral collection and acclimation
Specimens of the gorgonian coral S. suberosa (Pallas,
1766) were collected from coastal areas of Ho-Pin-Daw
Park (Peace Island Inlet, northern Keelung, Taiwan)
(25°090 4000 N, 121°450 2300 E). Colonies of S. suberosa were
cut with scissors at about 12–16 cm from the terminal tip
by SCUBA divers at water depths of 8–18 m. Specimens
were rapidly transferred in aerated plastic containers to the
laboratory, placed in quartz glass aquaria, and provided
with fresh seawater and air-supply, within 1 h after col-
lection. S. suberosa colonies were kept in a continuously
flowing seawater environment and were acclimated for
3 days to determine colony health before transfer to the
experimental containers. After showing tentacle extension
and feeding activity in the polyps, healthy colonies were
selected and used for the experiment. Seawater quality was
maintained at salinity ranges of 34.5–35.0 PSU, at
24–26°C, and 7–9 cm/s seawater flow speed, with an
illumination photoperiod from 0600 to 1800 hours at nat-
ural day light, and 1800 to 0600 hours in darkness. Fresh
Artemia franciscana nauplii were provided twice daily as
food (in the morning at 1000 hours and at night at
2300 hours). At the same time intervals, the behavior of the
corals and their polyp activities was observed daily for a
period of 3 days. The coral polyps of the three replicates
belonged to different colonies. We retransplanted the coral
colonies after experimentation back to the sampling area.
Sediment collection and handling
Red soil collected from a construction site near Keelung on
the northeast coast of Taiwan served as the terrestrial
sediment. Ocean sediment was collected by SCUBA divers
from the surface of the seabed by scooping around the
S. suberosa habitat at a depth of 15–20 m. The sediment
was brought to the laboratory in coolers. In the laboratory,
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Mar Biol (2011) 158:1301–1310
the sediment was washed with distilled water and dried at
140°C overnight, then ground in a mortar. We sieved and
selected the fine grain fraction (less than 105 lm) for the
suspension experiments for both sediment types. This fine
grain fraction typically occurs in the runoff water after
heavy rains.
Sedimentation experiment
Suspensions of terrestrial and oceanic sediments were
mixed in seawater. Suspensions of three sediment con-
centration levels, 50, 150 and 250 mg per liter seawater,
were used in the experiments. These sediment concentra-
tions represent a range of low, medium, and high concen-
trations and were used to understand the behavioral
changes that occur at various sediment concentrations.
A control group was exposed to seawater, without the
addition of sediment. Thus, there were seven different
treatments in the experiment, corresponding to two sedi-
ment sources, three concentration levels, and a control
group without sediment. Treatments were haphazardly
administered in seven glass aquaria, which contained nine
colonies of S. suberosa each.
All treatment groups and the control were provided with
an underwater pump (Supreme, Model: Ovation-200,
110–115 V, *60 Hz) and constant moderate air-supply in
the experimental aquaria to keep sediment in suspension.
The speed of flow was set at 7–9 cm s-1 which provided a
hydrodynamic condition where all sediment was kept in
suspension.
The experiments lasted for 15 days and each was
repeated three times. The three replicates were completed
within 1 week during each of the months of January,
March, and April. Since all experiments were conducted in
the laboratory, the environmental conditions (such as light,
temperature, salinity, and flow rate) were maintained
the same. Laboratory conditions were the same as those
during the acclimation period. During the experiment,
the polyp expansion activity of S. suberosa polyps was
recorded daily at 1000 hours (morning), 1630 hours
(evening), and 2300 hours (night) for each individual
polyp-colony, using a 5-point scale. Five ranking groups
(1, 2, 3, 4 and 5) of polyp expansion activity were cate-
gorized according to the percentage as 0, 1–25, 26–50,
51–75 and 76–100% of each colony, respectively. Mucus
sheet formation (MSF) and the number of colonies were
recorded once at 2300 hours every day. The recording of
polyp activity was possible even under heavy sedimenta-
tion environments because the polyp color was white in
contrast to the grayish color caused by the adhering sed-
iments. Mucus sheets were collected and fixed in 10%
formalin solution, and inspected for enmeshed particles by
light-microscopy.
Mar Biol (2011) 158:1301–1310
Record and logit transformation of the polyp activity
index and MSF
We used percentages to group and rank active polyps of each
colony since the colonies of S. suberosa were formed from
many individual polyps and the number of polyps in each
colony was not easy to count. The color of polyps was white
as distinguished from the coenosarc tissue which is light
brown in S. suberosa. The usage of percentages to estimate
the activity of polyps turned out to be a reliable character.
We designed the average ranked values for logit transformed
activity behavior (e.g. 12.5% would be the average value for
rank 0–25%). Recorded activity values were transformed for
each time point into an activity index as follows:
1 replicates, the group effect and concentration effect and
Activity index ¼ ðn1  0% þ n2  12:5% þ n3
9
37:5% þ n4  62:5% þ n5  87:5%Þ;
ð1Þ
where ni is the number of colonies (out of 9) with a
recorded value i. The index in (1) summarizes the average
percentage of colonies, according to their activities. The
larger the activity index (in percentage) is, the more active
the polyps of the respective colony were. We further
converted the daily activity index (averaged over three
indices within each day) for each treatment into the relative
odds at log scale (called log odds ratio or logit) by using
the following transformation:
 
Dailyactivity index
logit ðDailyactivity indexÞ ¼ log
1  Dailyactivity index
ð2Þ
The percentage of active polyps within a colony is thus
provided as the odds ratio of active polyps versus inactive
polyps.
The soft coral S. suberosa performs expanding behavior
to slough its mucus sheet. Removal of mucus also removes
the sediment grains. In the present study, quantification
applied to counting the number of colonies with mucus.
MSF was considered the number of coral colonies with
mucus at a particular monitoring time point. In the present
study MSF was scored by counting the number of coral
colonies with mucus at a particular monitoring time point.
The logit transformation was considered for the frequen-
cies (out of 9) of the MSF of coral colonies. If the fre-
quency was exactly zero or 100%, the transformation was
modified as log (0.5/8.5) or log (8.5/0.5), respectively.
Analysis of index data
We first summarized the sample mean and sample standard
deviation (SD) for the seven treatment groups, the results
1303
were obtained by pooling observations over time. The
95% confidence interval for each treatment group
mean was constructed as ðtreatment group sample meanÞ
1:96 ðtreatment group sample SDÞ=45.
The sample mean and standard deviation for the activity
index of each day of the experiment were computed for the
control and for the two sediment groups and averaged over
the replicates for the different concentrations. We con-
structed the 95% confidence intervals for the changes in
activity index over time accordingly for the control group,
the terrestrial group, and the oceanic sediment group.
We used ANOVA to test for the effects of sediment
source, concentration, and time which affected the varia-
tions of the activity index. Since our data consisted of three
their interactions were tested by comparing the mean
squares for each factor to the replicate error term. The time
effect and the other interaction terms were tested by
comparing the mean squares to the residual error. We also
examined the concentration and time effects more closely
by treating them as continuous variables.
Statistical analysis was carried out by a linear regression
method, using statistical software SPSS v.13 package.
Factors contributing to variation of the activity index and
the behavior of coral MSF were identified using the F test
(Zar 1999) at the significance level 0.05.
Results
;
Coral behavior
The behavior of corals under sedimentation revealed three
types of polyp activity patterns: withdrawal of polyps in the
coenosarc tissue (a), protrusion of polyps from the coe-
nosarc without extending their tentacles (b), and the normal
type of polyp body protrusion from the coenosarc tissue
with extension of their tentacles (c). In case (c) polyps
caught Artemia franciscana nauplii prey.
Mucus sheet formation and the physical composition
of mucus sheets
During corals produced mucus that formed a mucus sheet
covering the whole colonial body in the experimental
period. Under different sedimentation conditions, sediment
particles that adhered to this mucus sheet were indicated by
color changes. The color of mucus sheets mainly depended
on the type and amount of sediment entrapped in the
mucus. Higher loads of suspended materials induced a
higher production of mucus that in turn bound more sedi-
ment than lower sediment concentrations. In the control
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1304 Mar Biol (2011) 158:1301–1310

Fig. 1 Relationship between

activity index, frequency of
mucus sheet formation with
three concentrations for the two
sediment groups. The box
indicates the major distributed
region (75%) for the data, in
which the white line indicates
the median of the data, and the
upper and lower values of the
box indicate the 75th and 25th
quartiles of the data
respectively. There is a pair of
parentheses indicating the
boundary for outliers and each
black line indicate the extreme
value

group, corals produced less mucus and did so at a slower
pace than in the treatment groups (Fig. 1).
Mucus is colorless and transparent when observed under
the microscope. The mucus sheet affected coral feeding
behavior by hampering tentacle ability to capture nauplii of
the brine-shrimp Artemia franciscana.
Light microscopy revealed the complex composition of
mucus sheets which formed under sedimentation condi-
tions, showing tissue spicules, fungi, bacteria, and nema-
todes. Spicules that were emitted and subsequently
entrapped in the mucus provided evidence of injury to the
coral surface tissue. The spicules were wrapped in mucus-
sediment sheets when the corals were exposed to sus-
pended sediments for long periods of time.
Statistical results of activity behavior and MSF in study
The soft coral S. suberosa showed decreased activity of
polyps and increased MSF after exposure to suspended
sediments (Fig. 1). Both, the MSF frequency and activity
index significantly correlated with sediment concentration
in a linear fashion (Table 1). Higher sediment concentra-
tions correlated with more frequent formation of mucus
sheets (correlation, r = 0.613, p \ 0.0001) and a decrease
of activity of polyps (correlation, r = -0.293, p \ 0.0001).
The activity indices of polyps were negatively correlated
with the formation of mucus sheets (correlation = -0.536,
p = 0.048).
The analyses showed that the environments with dif-
ferent sediment sources revealed significantly different
activities (p \ 0.0001). The activity of polyps was reduced
with increasing exposure to suspended sediments as
compared to the control. Concentration effects showed
only a minor, not significant impact on activity of polyps
during sediment treatments (p = 0.0554). ANOVA was
repeated, treating time effect as a continuous variable.
Significant changes in activity frequencies were found over
time for each treatment group (p \ 0.0001, Fig. 2).
Plots for MSF based on transformed data over time for
polyp activities are shown in Fig. 3. The frequency of MSF
increased over time under all treatment conditions. The
change in this trend was faster with exposure to terrestrial
sediments than to oceanic sediments (p = 0.0001).
Table 2 shows the ANOVA results for the frequency
of MSF, which revealed that the sediment source
(p \ 0.0001), concentration (p \ 0.0001), and linear time
effects (p \ 0.0001) were all important factors explaining
the variations of mucus formation. In addition, the inter-
action between sediment source and time was significant
(p = 0.0001), indicating that the pattern changes in MSF
over time were different for the different sediment sources.
Models for trends in activity index and mucus sheet
formation
Model for the activity index
The sources of sediment (terrestrial or oceanic) were
treated as categorical factors; the concentrations (50, 150,
and 250 mg/L) and time index (t) were treated as contin-
uous variables. A mixed effect model was further fitted to
the activity index data and the fitted models for each
treatment group are shown in Table 3, in which all of the
effects were significant (p \ 0.05) except the time effect

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Table 1 Linear correlations (standard errors) of activity frequency of polyps, activity index, mucus sheet formation (MSF), and corresponding
sediment concentration
Linear correlations Activity frequency of polyps
Activity index 0.997 (0.004)
Mucus sheet formation -0.545 (0.047)
Sediment concentration -0.293 (\0.0001)
All of the correlations are significant at a significance level of p \ 0.05
for the control group (estimated coefficient = -0.0213,
p = 0.1708).
For the treatment groups, the time effects were impor-
tant and showed linear trends. Comparison of the estimated
slopes -0.0739 (terrestrial sediment), t statistic = -8.21,
p \ 0.0001) and -0.0662 (oceanic sediment), t statis-
tic = -4.14, p \ 0.0001) shows that the activity frequency
of polyps for terrestrial sediments decreased faster than for
oceanic sediments. The sedimentation stress on S. suberosa
was measurable only when the coral was exposed to the
silty sediment fraction (p = 0.0479, One-Way ANOVA),
whereas the sandy fraction of the two sediment types
did not affect corals during the experimental period
(p = 0.3368, One-Way ANOVA).
The inversion of the logit transformation was applied to
determine the predicted activity indices for low and high
concentrations for up to 50 days for the two sources of
sediments (Fig. 4). This shows an exponential decrease of
the expected activity index with time.
Model for the frequency of mucus sheet formation
Similar to the activity index, we fitted a mixed effect model
for transformed MSF data, and the time effects were
summarized by a quadratic relationship. The fitted models
for the different groups are shown in Table 4, where all of
the coefficients were significant. The predicted frequencies
of MSFs obtained by applying the inversion of the logit
transformation for the three sediment concentrations over
15 days for the two sources of sediments are shown in
Fig. 5. The corals exposed to terrestrial sediments pro-
duced significantly more mucus than the corals exposed to
oceanic sediments. Moreover, MSF occurred more fre-
quently at higher sediment concentrations. MSF frequency
reached 100% towards the end of the experiment, irre-
spective of the sediment concentration.
Discussion
Our study showed that for both sources of sediments, the
activity index decreased as the concentration of sediment
increased with the terrestrial sediment having a more
1305
Activity index Mucus sheet formation
-0.536 (0.048)
-0.300 (0.054) 0.613 (\0.0001)
deleterious effect. A previous study on the scleractinian
coral Montipora peltiformis showed that the stress level
after a short-term sedimentation episode increased linearly
with increasing amount and duration of sediment exposure
(Philipp and Fabricius 2003). In their study, M. peltiformis
was exposed to an average of 151.37 mg dry weight cm-2
of fine muddy coastal sediments for up to 36 h. We dem-
onstrated that such a linear relationship also existed at
lower and probably more environmentally representative
concentrations.
One important mechanism that minimizes sedimentation
damage is sediment removal by the coral polyps. Sediment
rejection behavior occurred when corals were exposed to
high siltation levels (Hubbard and Pocock 1972). Ciliary
currents, tentacle activities, body expansion and mucus
secretion behavior enmeshes sediment particles (Stafford-
Smith and Ormond 1992).
During expansion behavior corals fill their coenosarc
tissue up with body fluid. The body, therefore, appears lar-
ger than in untreated seawater. In our experiments we found
expansion behavior among the treatment groups not to differ
significantly from that of the control. It is therefore possible
that expansion behavior is a common activity among gor-
gonians. Stafford-Smith and Ormond (1992) reported that
almost all 42 coral species they investigated responded to
four different grain sizes by expansion behavior. Expansion
behavior, however, does not appear to be a good biomarker
for environmental stress caused by sedimentation. We found
frequency of expansion as a suitable indicator to the monitor
effects of sedimentation stress.
Weber et al. (2006) showed for Montipora peltiformis
that sediment removal rates depended on sediment prop-
erties; with sandy sediments removed more efficiently than
silty sediments. These authors also found that sandy grain
size fractions were rejected about three to four times more
effectively than nutrient-rich fine sediments, possibly due
to the greater volume and stickiness of the latter (measured
as settling rate, settling volume, and compaction). A neg-
ative relation between rejection efficiency and sediment
stickiness was shown by Fabricius and Wolanski (2000). In
their study, coral-inhabiting barnacles and their coral host,
Acropora sp., rejected nutrient-poor offshore sediments,
but not nutrient-rich, near-shore sediments.
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1306 Mar Biol (2011) 158:1301–1310

1.0
1.5
(a) 50 mg l-1
(a) 50 mg l-1
0.5 1.0

0.5
Activity index

MSF index
0.0

0.0

-0.5
-0.5

-1.0 Control Control

-1.0
Oceanic Oceanic
Terrestrial Terrestrial
-1.5 -1.5

1.0
1.5
(b) 150 mg l-1 (b) 150 mg l-1

0.5 1.0

0.5
Activity index

MSF index
0.0

0.0

-0.5
-0.5

-1.0
-1.0

-1.5 -1.5

1.0
1.5
(c) 250 mg l-1 (c) 250 mg l-1

0.5 1.0

0.5
Activity index

MSF index

0.0

0.0

-0.5
-0.5

-1.0
-1.0

-1.5 -1.5
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15

Time (Day) Time (Day)

Fig. 2 Patterns of the logit of activity indices over time for different Fig. 3 Patterns of mucus sheet formation (MSF) indices over time for
treatment groups different treatment groups

Previous studies demonstrated that coral rejection
behavior of sediments includes enhanced ciliary and ten-
tacle activity, colonies expansion, and mucus secretion
(Goldberg 2004; Levy et al. 2001). This was previously
demonstrated for mucus secretion in 42 species of Aus-
tralian scleractinian corals by Stafford-Smith and Ormond
(1992). Particular behaviors such as mesentery extrusion
and tissue pulsing have been shown for some species not
immediately related to S. suberosa. We demonstrated that
colony expansion could help S. suberosa to break the
mucus sheet, which is subsequently carried away by water
currents. Hermatypic corals exposed to suspended sedi-
ment show mucus formation similar to the gorgonian
S. suberosa. The production of mucus provides a substantial

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Mar Biol (2011) 158:1301–1310 1307

Table 2 ANOVA results for the frequency of mucus sheet formation

(MSF) of S. suberosa for different sediment suspension treatments
(control group, ocean sediment and land sediment groups; conc.
concentration)
Source of df Sum of Mean F value p value
variations squares squares

Sediment 2 366.75 183.37 245.41 \0.0001

Concentration 2 86.13 43.06 57.63 \0.0001
Time 1 131.00 131.00 175.32 \0.0001
Sediment 9 conc. 2 0.99 0.49 0.66 0.5171
Sediment 9 time 2 20.72 10.36 13.87 \0.0001
Conc. 9 time 2 10.21 5.11 7.11 0.0010

Table 3 Fitted mixed effect model for transformed activity indices

(t time, conc. concentration) of S. suberosa for different sediment
suspension treatments Fig. 5 The predicted the frequency of mucus sheet formation over
time for low and high concentrations of two sediment groups
Treatment groups Fitted model

Control y^ ¼ 0:4602  0:0213t sink of material by S. suberosa under sedimentary

ð0:174Þ ð0:016Þ

Terrestrial sediments y^ ¼ 0:4896  0:0739t  0:0013 ðconc:Þ conditions.

ð0:136Þ ð0:009Þ ð0:0006Þ Sediments exert profound influence on the energetic
Oceanic sediments y^ ¼ 0:5887  0:0662t  0:0013 ðconc:Þ balance of symbiotic Scleractinia and Alcyonacea. This is
ð0:136Þ ð0:016Þ ð0:0006Þ
partly due to absorption or reflection of light by the sedi-
The numbers in the parentheses are the standard errors of the esti- ment, but can also be caused by stimulation of mucus
mated coefficients shown above
production. Both of these factors shift the energy balance
of corals because of decreased photosynthesis by algae and
increased loss of mucus. Under heavy sedimentation con-
ditions, corals shut down most of their normal metabolic
functions in favor of producing large quantities of mucus,
which aid sediment removal (Stafford-Smith and Ormond
1992). The practice of producing mucus sheets is a costly
way of reacting to the environment, as in most cases,
carbon equivalents of several days of production are
released (Coffroth 1980). Over long periods of time, this
creates severe problems for corals and may very well
explain long-term physiological changes as reported by
Philipp and Fabricius (2003), although these authors
worked on horizontal flat coral surfaces whereas S. sube-
rosa is an erect branching species.
Sediments affect corals in several ways, being suspended
Fig. 4 The predicted the frequency of polyp active over time for low
and high concentrations of two sediment groups or becoming settled on surfaces of corals (Fabricius et al.

Table 4 Fitted mixed effect model for the transformed mucus sheet formation frequency (MSF) (t time, conc. concentration, (conc. 9 t)
interaction between conc. and time) of S. suberosa for different sediment suspension treatments
Treatment group Fitted model

Control y^ ¼  2:0938 þ 0:2417t  0:0120t2

ð0:358Þ ð0:047Þ ð0:003Þ

Terrestrial sediments y^ ¼  1:0327 þ 0:0111ðconc:Þ þ 0:3938t  0:0120t2  0:0005ðconc:  tÞ

ð0:341Þ ð0:002Þ ð0:046Þ ð0:003Þ ð0:0001Þ

Oceanic sediments y^ ¼  2:4021 þ 0:0111ðconc:Þ þ 0:4870t  0:0120t2  0:0005ðconc:  tÞ

ð0:341Þ ð0:002Þ ð0:046Þ ð0:003Þ ð0:0001Þ

The numbers in parentheses represent the standard errors of the estimated coefficients shown above

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1308
2003a, b; Fabricius 2005). Corals exposed to high sediment
concentrations may have high sediment tolerance (Sofonia
and Anthony 2008) or typically show sub-lethal (Rice and
Hunter 1992) or lethal effects (Hoitink and Hoekstra 2003).
In addition, mucus secretion from coral epithelia increases
carbon loss (Rogers 1990; Stafford-Smith and Ormond
1992). Sediment rejection efficiency varies among coral
species and also among sediment types (Stafford-Smith
1993; Levy et al. 2006). Sedimentation stress and recovery
after short-term sedimentation depend on the amount and
duration of sediment deposition on the corals (Philipp and
Fabricius 2003). Sedimentation inhibits the settlement of
coral larvae and smothers newly settled juveniles (Birrell
et al. 2005). Some reports even suggest that sedimentation
causes coral death (Rogers 1990; Rice and Hunter 1992). At
the same time they also increase the metabolic activity
and respiration of corals (Stafford-Smith 1993; Riegl and
Branch 1995; Philipp and Fabricius 2003). Hubbard and
Pocock (1972) showed that sediment rejecting activities by
polyps, such as tentacle movement, tissue swelling, and
ciliary activity increase energy consumption (Riegl and
Branch 1995) that otherwise could be allocated to processes
such as growth and reproduction (Logan 1988; Stafford-
Smith and Ormond 1992; Riegl and Branch 1995; Philipp
and Fabricius 2003). Respiratory rate due to a high siltation
environment may be 30% higher compared to clear water
conditions (Riegl and Branch 1995). High microbial den-
sities on resuspended and settled particles can quickly lead
to oxygen depletion, followed by sulfate oxidation and
hydrogen sulfide development in sediments (Schulz and
Zabel 2000). Reduced circulation reduces oxygen
exchange, and further promotes anoxia and hydrogen sul-
fide formation (Boudreau and Jørgensen 2001). It remains
to be shown whether resulting anoxia and microbially
mediated development of the cell toxin hydrogen sulfide are
the primary causes of coral poisoning. Sedimentation not
only interferes with the coral’s energy balance by enhancing
respiratory losses and reducing photosynthetic production,
but also reduces heterotrophic energy gain by interfering
with the prey-capturing apparatus. This was demonstrated
for S. suberosa by Dai and Lin (1993).
Sedimentation is a major cause of coral mortality, par-
ticularly in the initial life stages (Nugues and Roberts
2003). It can locally reduce recruitment rates, and affect a
range of life history parameters in adult corals and can
enhance diseases (Richardson 1998). Nugues and Roberts
(2003) showed that partial mortality and fission in colonies
of four common massive coral species depended on the
sites that were exposed differently to river sediments in St.
Lucia, West Indies. Studies on the effect of sedimentation
frequently focus on the amount (weight) of sediment, while
the fact that sediment consists of a vast and contrasting
array of mineral particles of various grain sizes, organic
123
Mar Biol (2011) 158:1301–1310
particles, and living organisms, detritus, mucus, and exo-
polymeric particles remains largely unstudied (Fabricius
2005). Sediment can also be a carrier of adsorbed or par-
ticulate nutrients and contaminants. With the exception of
grain size, not much is known about differences in sedi-
ment characteristics that may influence the survival of
corals exposed to sedimentation (Fabricius et al. 2003a, b).
Low-level sedimentation of sticky, muddy marine snow
aggregates affect small reef organisms, such as epibiotic
barnacles on corals, more than intermediate amounts of
mineral sediments (Fabricius and Wolanski 2000). It is
likely that sediment characteristics, such as the tendency to
form sticky marine snow aggregates and differences in
organic and microbial concentrations, also alter the effect
of sedimentation on coral recruits.
Sedimentation also leads to coral bleaching and epi-
thelial necrosis (Rice and Hunter 1992; Pain 1996). During
bleaching, symbiotic algae are expelled from coral tissues,
die, or lose their photosynthetic pigments (Anthony et al.
2007). Suspended materials such as sediments decrease
light penetration and reduce the photosynthetic capability
of coral symbionts (Dallmeyer et al. 1982; Harrington et al.
2005). Sedimentation results in an energetic cost for pho-
tosynthetic organisms through damage of their photosyn-
thetic apparatus, or increasing their metabolism due to
damage and avoidance behavior or molecular repair
mechanisms (Hoogenboom et al. 2006). Previous studies
found a decline of up to 22% of symbiotic algal densities
after sedimentation events (Dallmeyer et al. 1982). After
such disturbances, recovery can occur, but this can take
6–10 weeks, if the corals survived the event (Jaap 1979).
Several studies described the growth and behavioral
reactions of Scleractinia (stony corals) to sedimentation
(Stafford-Smith 1993; McClanahan and Obura 1997;
Fabricius et al. 2003a, b). Coral activity in a high siltation
environment reduces the net production of corals
(Dallmeyer et al. 1982). Most sedimentation studies have
been concerned with scleractinian corals. The very same
gorgonian species studied here were otherwise only studied
by Dai and Lin (1993). The latter authors showed that
S. suberosa was affected by sedimentation due to reduced
heterotrophic energy gain caused by interference with the
prey-capturing apparatus.
Accurate prediction of sedimentation effects at a given
site is an important task for coral reef risk assessments, as it
is difficult to correlate the rate of sedimentation with
damage (Wessling et al. 1999). Rogers (1990) reviewed the
sedimentation effect on coral ecosystem and found that low
concentration of sediment cause detrimental effects; vice
versa may high load of sediments not show serious dam-
age. Brown et al. (1990) reported that rapid recovery is
possible even after intense sediment damage. Several
studies showed that confounding anthropogenic influences
Mar Biol (2011) 158:1301–1310
provide interactive effects from fishing (Birkeland 1997),
from nutrient enrichment (Ferrier-Pages et al. 2000), and
from physical natural disturbances such as hurricanes or
monsoons (Wilkinson 2004).
Riegl and Branch (1995) found that sedimentation
affected coral metabolism and decreased the photosyn-
thetic production. They reported that carbon-loss by mucus
secretion shifted the energy balance of corals. Mucus
production of corals in sedimentary environments rose by
65% in comparison to unaffected corals, which is an
average 35% higher than the daily respiration (Riegl and
Branch 1995). We found that MSF was affected by all
sediment concentrations. MSF increased with increased
environmental sediment concentrations. We found that the
frequency of MSF in S. suberosa was higher when it was
exposed to terrestrial than to oceanic sediments. A previous
study by Yamasu and Mizofuchi (1989) also found that
terrestrial soil sediments increased the respiration rate of
the coral Fungia repanda.
Synergistic effects between sedimentation stress and
nutrient concentrations in the sediment have been reported
for corals (Fabricius et al. 2003a, b). Similarly, synergistic
effects between sedimentation and herbicides are found in
crustose coralline algae, where sedimentation stress is
greatly enhanced by the presence of traces of the herbicide
diuron (Harrington et al. 2005). Further investigation of
synergistic effects between various sediment properties is
needed to understand the potential harm to corals from
exposure to sedimentation.
Acknowledgments We acknowledge our debt of gratitude for the
assistance provided by the technicians from J. S. Hwang’s laboratory
for their contribution to the field-sampling program. We are grateful
to the National Science Council of Taiwan, ROC (NSC grant No.
97-2621-B-019-004) and the Center of Excellence for Marine Bio-
environment and Biotechnology of National Taiwan Ocean Univer-
sity (99529001A) for financial support to J.S. Hwang.
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