Proceedings 9"' International Coral Reef Symposium. Bali, Indonesia 23-27 October 2000. Vol.

Effects of sedimentation on coral settlement and survivorship

R. Babcock ' and L. Smith' .
ABSTRACT
Settlement and survival of juveniles are key components of recruitment, which is in turn crucial to the persistence of
coral populations. Settlement and juvenile survival of the coral Acropora millepora were measured in situ using
experimentally elevated sedimentation rates. Sedimentation rates ranged between 0.76-1.32 mg.cm"=d'' for reference
sites and 1.88-11.70 mg.cm' a .d'' in sediment treatments. Settlement was significantly lower in the sediment treatments
where it reached only 71% of levels in reference areas. The number of juvenile corals surviving at sediment treated sites
after 8 months was 39% of levels at reference sites, however mortality did not vary significantly between treatments.
These effects indicate that even relatively small increases in sediment accumulation would have negative impacts on
coral recruitment.

Keywords Sediment, Corals, Acropora millepora, Recruit-
ment, Settlement
Introduction
Sediment and turbidity are recognised as important
ecological correlates of the distribution of reef building corals
(Stoddart 1969, Rogers 1990). Nevertheless there are many
gaps in our understanding of the mechanisms through which
sediment may affect corals ecology. Increasing levels of
human disturbance in coastal catchments and the
transportation of sediment into nearshore coastal waters and
onto reefs has increased the need to understand the precise
role of sedimentation on reef corals. While thriving reefs are
found in turbid conditions (Bull 1982) their coral
assemblages tend to differ in composition from those on reefs
subjected to lower levels of turbidity and sedimentation.
Turbid reefs are often dominated by massive forms, while
those in clearer waters are characterised by branching forms,
particularly acroporids (Done 1982).
Sedimentation and suspended sediment in the water
column may affect coral populations and community
structure by smothering adult corals or imposing
physiological costs by reducing light availability for
photosynthesis or increasing the need for active sediment
removal (Reigl and Branch 1995). Understanding of
sedimentation effects is complicated by interspecific
differences in morphology and sediment rejection ability
(Stafford Smith 1993), variations in the ability of corals to
utilise heterotrophic modes of nutrition (Anthony 2000), and
the fact that high sedimentation rates tend to co-vary with
other environmental and biological factors such as wave
exposure and larval supply (Done 1982, Sammarco 1991).
A further way in which sediment may affect coral
community structure is by influencing recruitment rates, and
the small size of juvenile corals highlights their potential
susceptibility. Coral recruitment has been correlated with
environmental parameters including suspended sediment load
(Hunte and Wittenberg 1992, Wittenberg and Hunte 1992)
however, since these studies utilize environmental
gradients, this could be the result of several co-varying
factors such as salinity, nutrients, algal cover, light
penetration or larval supply. In order to eliminate these
confounding factors, field-based artificial manipulations of
sedimentation rates have been used to measure sediment
effects on both larval settlement and subsequent survival
of the staghorn coral Acropora millepora.
Methods
Larvae of Acropora millepora were reared by
conventional procedures (Babcock and Heyward 1986),
using eggs spawned on the evening of March 19' h at Coral
Bay (23° 09' 5; 113° 45' E) Ningaloo Reef Tract, Western
Australia in March 1998. The extensive Ningaloo reefs
are part barrier and part fringing reefs characterised by
recent coral structures overlying Pleistocene substrate
( Veron and Marsh 1988). An extensive shallow lagoon 2-
l2 m in depth is protected from oceanic swells. Climate in
the region is arid and the reefs are subject to naturally low
levels of terrestrial runoff and sedimentation (visibility in
the lagoon is > 10 m). Experimental sedimentation rates
were manipulated using rammed earth bricks which
softened after submersion allowing the wave action at the
site to distribute sediment in their immediate vicinity. The
bricks (300 x 300 x 150 mm, 17 kg) were composed of
terrestrial sediment of no less than 90% silt/clay (without
cement stabiliser). Five bricks were deployed within each
of two 6.25 m ' treatment sites, approximately 200 m
offshore from Coral Bay at a depth of approximately 4 m.
Additional bricks were added periodically to ensure
continued sediment supply. The first deployment was on
March 22 ntl , followed by deployments on March 28 and
thereafter on May 7 h and June 10 th . Treatment sites were
separated by approximately 50 m.
Sedimentation rates were measured in the treatments
and in their paired reference areas located some 15 m
away, using low-profile sediment collection devices

' University of Auckland Leigh Marine Loboimory. PO Box 349 Warkworth, New Zealand, r.habcock@i auekland.ac.nz.
1 Australian Institute of Marine Science. P013OX 264 Dampier Western Australia. 6713. Australia

245
consisting of a 3 mm thick perspex plate to which a
TM
scotchbrite scrubber pad (100 x 150 mm) was attached
using velcro strips. This assembly was attached to the reef in
a horizontal position. The structure of the scrubber pad
resembles algal turf and was no more than 10 mm above the
level of the surrounding turf, providing for near-bed trapping
of locally re-suspended sediment particles. Four sediment
collector pads were deployed at each site.
Scrubber pads were deployed on March 21 `' and
sedimentation rates were sampled by collecting and replacing
the scrubber pads on March 27' , May 76 and June 10 '" . Both
the mount and the scrubber were removed from the
substratum, taking care not to lose any sediment. Both were
placed in a zip-lock bag in-situ, the scrubber released, and
both shaken for approximately 20 sec. until all sediment had
fallen into the bag. The scrubber and mount were then
removed and the bag sealed. Excess seawater was removed
onshore using filter paper and a vacuum pump, and the
sediment samples were then placed in an oven at 80 °C until
stable and weighed.
Larval settlement was manipulated using enclosures
consisting of transparent hemispherical bowls 12 cm in
diameter. Water was able to circulate through the enclosures
via two rectangular mesh covered (200 m) openings on
opposing sides of the bowl [see Babcock and Mundy (1996)
for details of methods]. Coral settlement was measured using
'
square terracotta tiles (120 x 120 x 10 mm). A hole was
drilled in the centre of each tile, which was fixed to the reef
matnx in a horizontal position using a stainless steel bolt
hammered into the substratum. A 10 mm plastic spacer
maintained a space between the tile and the substratum.
Sixteen tiles were deployed at each site. Plates were placed
in the sea for conditioning on March 12 'h , and were fixed to
the reef on March 21'. The enclosures were placed over the
tiles on March 26, four days after the start of sediment
treatments. Approximately 1200 ±100 six-day old larvae of
A. millepora were then injected into the enclosures. Tiles
were examined 4 days after the introduction of larvae to
determine the number of settlers. While the enclosures
would have reduced sediment accumulation during this time,
there was an obvious accumulation of sediment on the upper
surfaces of the tiles, as well as some adhering to the vertical
edges of the tiles. During the census corals on each tile were
mapped and then returned to their previous point of
attachment. All juveniles were recorded as dead or alive
at the subsequent censuses (May 7ih , June 10'h and
November 20'5. The tiles were always immersed in
seawater during measurement.
Due to non-normality of the data (many settlers on a
few plates, few settlers/survivors on most plates) their
distribution was closer to Poisson or binomial than
normal. Accordingly, settlement and survival data were
modeled under the Poisson and binomial distributions
respectively (SAS procedure GENMOD). Values
presented for test statistics were obtained after backward
fitting of saturated models, where non-significant effects
were progressively - removed. Analysis of variance
(Factors Treatment and Time fixed; Factor Site random
and nested within Treatment and Time) was used for the
sedimentation rate data, which was normally distributed.
Results
Sedimentation rates
By November 1 998 the scrubbers had been removed
(probably by grazing fishes) or had become completely
overgrown by the alga Dic tyota. Consequently the
sediment loads were not measured in November 1 998.
Even the June samples had a large standing crop of algae
on the scrubbers which is likely to have effected their
efficiency. One scrubber was lost at this time.
Sedimentation rates in the treatments ranged from 11.7
'
t5.0 to 1.88 t1.0 mg.cm°.day and were significantly
higher (F ), z=51, p<.001) than those in the reference areas
which ranged between 0.8 1-0.3 to 1.2 t0.2 mg.cm' .day '
(Table I). There was no significant variation between
sites, though over the course of the experiment
sedimentation rates decreased significantly at the
treatment sites (F ), r=4.8, p=0.02).
Settlement and survival
Settlement in the sediment treatment areas was 71% of
that in the reference areas (Table 2, x = 5.32, p=0.021).
2
Settlement varied significantly among plate surfaces, with
the majority of settlement taking place on the
undersurfaces of plates (Fig. I, Chi-squared = 277.9,
p < 0001, model: treatment aspect).

' '
Table I. Sedimentation rates at Ningaloo experimental sites. Sedimentation rates are mg.cm .day' ± 95% ci.
Reference sites Sediment treatment sites
March May June Overall March May June Overall
Site
7.93 t 1.9 2.44 t0.8 2.13 t0.5 4.17 t 1.7
I 1.01 ±0.2 0.91 ±0.5 1.24 t0.4 1.06±0.2
1.18±0.2 1.14 t0.3 15.47 t87 2.42±0.8 2.16 t0.5 7.10±4.7
2 1.62 ±0.3 0.62 t0.2
Total 1.32±0.3 0.76± 0.3 1.21t 0.2 1.10 t 0.2 11.70 t4.9 2.43 ±0.6 1.88 t0.7 5.34±2.4

246


Table Z . Sediment treatment and coral ' settlement . and survival . Summary of Log-linear analysis, values are, type 3
likelihood ratio statistic s

A . Settlement

Source Num DF Den DF r F Value P>F: Chi-Square - P > Chi-Square

Treatment " I' 92 - 532 = 0 .0234 i 53 2 0 .0211 '
x.
Aspect 2 92 138 .85 <0 .0001 277 .7 <0 .000 1

B . Corals surviving after 8 months

Source Num DF Den DF F Value P F Chi-Square . . P > Chi-Square
Treatment 1 91 fl, 8 .26 0.005 8 .26 0 .00 4
Site t, 1. 91 4 .14 i 0.045 4 .14 . 0 .041 8
Aspect 2 . . 91 . : 39 .82 <0 .0001 . 79 .64 <0 .0001

The survival of A . millepora to May, June an d

November was variable and did not differ significantl y
with treatment. Consequently ; by November numbers
alive in the sediment treatments remained significantl y
%lower -(Fig . 2,`Chi-squared ='7 .99; p=0 .0047, model :
treatment site aspect) though by this time they were 39 %
m
a25 of the number surviving in the reference sites . While there
a. was also significant variation in recruit numbers due t o
N
plate aspect and site effects, there were no significan t
,8 interactions amongst these factori or sediment treatment .
' Overall survival, from settlement to 8 months was 8 . 2
±1 .5% (sd) .
a

undersurfaces sides ' uppersurfaces total Discussio n

Fig 1 . Effects of sedimentation rate on , settlement of A . , .
Increased sedimentation rates can reduce rates of
millepora . Data for average settlement on each plate surface ,
and for total settlement per plate are from April, immediately settlement • in Acropore millepora . Reduced settlement
I for rates are 'likely to have implications for the recruitmen t
following settlement of, the ,larvae . See Table , into coral populations . The magnitude of sedimentatio n
sedimentation data . effects on settlemenbWere initially small, 71% of rates at
reference areas, but by - 8 months after settlement number s
eE s . • control of juvenile corals in the sediment treatments were down t o
0 sediment , . a" 39%' of those in reference areas . Sediment effects on
- settlement have been addressed previously by laborator y
d 2 manipulations with conflicting results, with both neutra l
a (Babcock and Davies 1991) and negative presence of
• • • elevated sediment levels . In natural populations subjecte d
3 t to increased sedimentation these differences in th e
numbers of settlers and juvenile corals are likely t o
translate into reductions in recruitment to adul t
o O populations . Reduced recruitment is in turn likely to have
significant effects on coral populations (Hughes and
undersurfaces sides uppersurfaces total Tanner 2000) .
The overall reduction in settlement and juvenile s
Fig 2 . Effects of sedimentation rate on recruitment of A . surviving to 8 months was in agreement with field studie s
millepora . Data for numbers of juveniles on each plate that have demonstrated a correlation betwee n
surface, and for total numbers per plate are from November, sedimentation or turbidity and coral recruitment rate s
approximately 8 months after settlement . . See Table 1 for (Wittenberg and Hunt 1992, Tomascik 1991) . Our stud y
sedimentation data. . indicates that even relatively small increases in
sedimentation may have significant effects on recruitment .

24 7
 Experimental sedimentation rates were not constant and
declined over the period of the experiment, a fact that could
be responsible for the similar rates of survival between
treatments. Given that the decline in sedimentation rate over
ti me was a product of a decline in the replacement frequency
of the mud-bricks, rather than an inherent property of the
methodology, there is clearly potential to extend the concept
of manipulating sedimentation rates in situ. Such
manipulations will allow us to better isolate sedimentation
from other environmental factors and determine its
importance as an environmental property affecting coral
population dynamics and coral reef community structure.
Similar methods have previously been applied successfully to
algal populations where increased sediment loads have been
shown to reduce recruitment rates in the brown alga
Sargassum microphyllum (Omar et al 1998).
Acknowledgments Thanks to Andrew Hayward, staff of
AIMS WA, also to T Willis for advice on log-linear models.
This research was supported by University of Auckland and
AIMS.
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