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Effects of suspended sediment on survival, growth, and nutritional
condition of green-lipped mussel spat (Perna canaliculus, Gmelin, 1791)
Brandy S. Biggar
a,*
, Andrew Jeffs
a,b
, Jenny R. Hillman
a
a
Institute of Marine Sciences, The University of Auckland, 23 Symonds Street, Auckland 1010, New Zealand
b
School of Biological Sciences, The University of Auckland, 3 Symonds Street, Auckland 1010, New Zealand
ARTICLE INFO
Dataset link: Effects of suspended sediment on
survival, growth, and nutritional condition of
green-lipped mussel spat (Perna canaliculus,
Gmelin, 1791) (Original data)
Keywords:
SS
Bivalve
Juvenile
Growth
Sediment
Turbidity
Laboratory trial
ABSTRACT
Suspended sediment is a signicant current threat to coastal ecosystems in many parts of the world. Sediment
runoff into the ocean is increasing due to numerous human activities, such as agriculture, deforestation, con-
struction, and mining. Additionally, climate change is impacting local weather patterns, with many regions
experiencing marked changes in the frequency and extent of precipitation. Extreme weather events, such as
cyclones, can increase suspended sediment (SS) in coastal waters by up to 100-fold. This rapid change in SS can
negatively affect lter-feeding bivalves by diminishing their feeding efciency, often preventing feeding until the
sediment settles. In New Zealand, juveniles of the green-lipped mussel (Perna canaliculus) have rudimentary
structures for capturing and sorting food particles from the seston, which are prone to clogging and damage. In
this study, the effects of a range of SS concentrations (0–1250 mg L
−1
) on the survival, growth, and nutritional
condition of juvenile green-lipped mussels (1–2 mm SL) were determined over two time scales (5 and 30 days) in
controlled laboratory experiments. Neither mortality nor nutritional condition were impacted by SS. However,
the presence of SS positively affected growth (p<0.05) and mussel settlement location (p<0.01) at both time
scales. The results show that, under these conditions, SS levels ≤1250 mg L
−1
are not apparently harmful to
P. canaliculus spat and may even be advantageous.
1. Introduction
Elevated suspended sediment (SS) in coastal waters is increasingly
recognised as a signicant ecological threat to aquatic species, especially
suspension feeders (Ellis et al., 2002;Lohrer et al., 2006;Lummer et al.,
2016). Suspended sediment in coastal waters originates from the erosion
and dispersal of terrestrial sediment and can be altered by several nat-
ural factors, including tectonic activity, lithology, precipitation, and
vegetation (Hicks et al., 1996, 2011;Trustrum et al., 1999). Moun-
tainous Southwest Pacic islands are particularly prone to erosion due
to their high ocean exposure, frequent tectonism and volcanism,
mountainous terrain, young and erodible rock, periodic intense rainfall
and weather events (e.g., cyclones, orographic induced rainfall), and
short drainage basins (Gayer et al., 2019;Goldsmith et al., 2008;Kao
and Liu, 2002;Milliman and Meade, 1983;Pariyar et al., 2020). These
islands have some of the highest global weathering and erosion rates,
collectively contributing >30 % of the global annual ocean sediment
yield (Carey et al., 2002;Hicks et al., 1996, 2011;Meyer et al., 2017;
Milliman et al., 1999). Human colonisation additionally exacerbates this
high erosion with urban development, agriculture, mining, and defor-
estation (McLeod et al., 2011;Milliman et al., 1999;Paul, 2012).
Many Southwest Pacic islands have been severely deforested
(Nunn, 1990;Rolette and Diamond, 2004). Easter Island, for example,
was entirely deforested by the prehistoric Rapanui civilisation (Rull,
2020; but see Davis et al., 2024), while substantial native vegetation has
also been cleared in New Zealand (Adams, 1979;Gomez et al., 2009;
Hicks et al., 2011; Rolett and Diamond, 2004; Rull, 2020;Selby, 1972).
Deforestation leads to slope instability and soil loss (Nunn, 1990),
increasing sediment runoff yields by up to 10 times (2–3 times on
average; Hicks et al., 2011;Milliman and Meade, 1983;Milliman and
Syvitski, 1992). Many Southwest Pacic islands are also highly vulner-
able to climate change, as increased frequency and intensity of rainfall,
oods, monsoons, and cyclones are all projected for their future (IPCC,
2023;Pariyar et al., 2020). Cyclones and intense precipitation can
trigger high erosion events, such as landslides and oods (Gayer et al.,
2019). For example, Typhoon Mindulle deposited 61 million t of sedi-
ment into the Choshui River in Taiwan (Goldsmith et al., 2008), while in
New Zealand, SS in estuaries typically increases from 10 to 1000 mg L
−1
* Corresponding author.
E-mail addresses: brandy.biggar@auckland.ac.nz (B.S. Biggar), a.jeffs@auckland.ac.nz (A. Jeffs), j.hillman@auckland.ac.nz (J.R. Hillman).
Contents lists available at ScienceDirect
Journal of Experimental Marine Biology and Ecology
journal homepage: www.elsevier.com/locate/jembe
https://doi.org/10.1016/j.jembe.2024.152074
Received 21 July 2024; Received in revised form 15 November 2024; Accepted 16 December 2024
Journal of Experimental Marine Biology and Ecology 582 (2025) 152074
0022-0981/© 2024 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license ( http://creativecommons.org/licenses/by/4.0/ ).
during large storms (Ellis et al., 2002;Hewitt and Norkko, 2007). Hence,
human activities substantially impact erosion, increasing sediment
runoff to the ocean.
Extreme sediment uxes threaten estuarine and coastal lter-feeding
organisms, such as bivalves, by rapidly and dramatically increasing SS
levels. This leaves organisms little time to respond, leading to burial,
suffocation, and starvation (Poirier et al., 2021). High SS levels pose a
signicant threat to lter-feeding bivalves (Sa et al., 2007), but the
level that triggers a response and the type of response depends on
numerous factors, including the bivalve species (Sa et al., 2007;Ward
and MacDonald, 1996), their adaptability (Lohrer et al., 2006;Tuttle-
Raycraft and Ackerman, 2019), exposure characteristics (Grant and
Thorpe, 1991;Hewitt and Norkko, 2007), and SS quality (Tuttle-Ray-
craft and Ackerman, 2018;Ward and MacDonald, 1996). Stressors like
SS may also have life-stage-specic impacts due to ontological differ-
ences in lifestyle (e.g., sessile/planktonic, benthic/pelagic) and feeding
(e.g., morphology, diet). For example, the responses of three different
freshwater mussels (Lampsilis fasciola, Lampsilis siliquoidea, and Sag-
ittunio nasuta) varied markedly by life stage when exposed to elevated
SS, with clearance rates increasing in the 1-week-old mussels and
decreasing in older juveniles and adults (Tuttle-Raycraft et al., 2017). It
is commonly recognised that younger life stages of marine invertebrates
are less tolerant of environmental stressors than adults (Chelyadina
et al., 2017;Ringwood, 1993). However, the life stage studies that
support these claims are less common and primarily relate to heavy
metal stressors and not SS (Calabrese et al., 1973;Connor, 1972;Mar-
kich, 2021;Martin et al., 1981;Ringwood, 1993). Meanwhile, studies
that are focused on SS have found variable results. For example, Pinna
nobilis and Mercenaria mercenaria juvenile growth (length and weight)
were negatively affected by SS (Acarli et al., 2011;Bricelj and Malouf,
1984), while Limnoperna fortunei larval growth was positively affected
(Eilers et al., 2011), and Corbicula uminea growth was unaffected (Foe
and Knight, 1985). Moreover, Crassostrea virginica juveniles were less
abundant in high turbidity (Reustle and Smee, 2020), and Dreissena
polymorpha veligers were more abundant in high turbidity (Barnard
et al., 2003); however, neither study characterised the suspended par-
ticles in the water column. Therefore, the specic impacts of increased
SS on bivalves vary and are challenging to predict.
Bivalve settlement and recruitment may also be negatively affected
by high SS concentrations, disrupting settlement cues (e.g., chemical
and light), reducing oxygen availability, or coating settlement sub-
strates, making them unfavourable or difcult to settle on (Poirier et al.,
2021). For example, C. virginica recruitment is negatively affected by
sediment stress (Thomsen and McGlathery, 2006) and high turbidity
(Reustle and Smee, 2020), and juvenile Margaritifera margaritifera
presence is correlated with turbidity and inorganic sedimentation
(¨
Osterling et al., 2010). Restoration of wild mussel populations in New
Zealand has been limited by recruitment, with no recruits observed in
over a year in South Island restoration projects (Benjamin et al., 2022,
2023;Toone et al., 2023) and only three recruiting individuals found
within 2 years in a North Island study (Wilcox et al., 2018). This lack of
recruitment suggests an unfavourable environment for recruitment,
with SS implicated as a likely reason due to dramatic increases in sedi-
ment discharge in these coastal regions (Handley et al., 2017;Swales
et al., 2016). In green-lipped mussel (Perna canaliculus) early juveniles,
the lter-feeding structures are not fully developed, have lower particle
capture efciency and a reduced ability to sort particles compared to
adults, and are prone to blockage and damage (Gui et al., 2016a, 2016b).
Elevated mortality, reduced growth, and lack of settlement could be
expected when juvenile green-lipped mussels are exposed to elevated SS
because of the increased energy requirements needed to lter sufcient
food particles due to reduced feeding efciency. For example, low green-
lipped mussel settlement near the seaoor has been associated with high
turbidity (Toone et al., 2023). To determine whether high SS is limiting
mussel recruitment in New Zealand, the current study explores the ef-
fects of a range of suspended sediment concentrations on the survival,
growth, and nutritional condition of P. canaliculus early juveniles.
2. Methods
2.1. Mussel collection and acclimatisation
The experiments in this study used hatchery-reared, green-lipped
mussel early juveniles, commonly known as spat. The spat were supplied
by a commercial mussel hatchery (SPATnz Ltd. Nelson, New Zealand) in
two batches (May 2023 and March 2024) as plantigrades of ~1–2 mm
mean shell length (SL), approximately one-month post-metamorphosis.
The hatchery utilises a continuous-ow seawater system at 20 ◦C, with
seawater ltered to 1
μ
m and treated with UV.
After transport to the seawater facilities at the University of Auck-
land’s School of Biological Sciences, mussel spat were added freely to
the water column of 20 L tanks in 18 ◦C ltered and sterilised seawater
(FSW) with constant aeration in a temperature-controlled laboratory.
2.2. Mussel husbandry
Before experimentation, mussels were acclimatised for 1 week.
During this period, FSW was replaced in 20 L holding tanks every second
day. Tanks were rinsed thoroughly with freshwater once per week to
reduce bacterial load; mussels were gently detached from tank walls
with a soft bottle brush, gently poured over a 200
μ
m sieve, rinsed, and
returned to the tank with FSW. Mussels were fed one species of
laboratory-cultured axenic microalgae (i.e., Tisochrysis lutea, Nanno-
chloris atomus, Chaetoceros muelleri, and Diacronema lutheri) in rotation at
~500,000 cells mussel
−1
day
−1
(concentration determined by Muse®
Cell Analyser, Millipore Sigma), which is sufcient to meet their nutri-
tional requirements to support growth (Sanjayasari and Jeffs, 2019).
2.3. Sediment collection and preparation
Surface benthic sediment was collected from a mudat in the inner
Whangateau Harbour, New Zealand, which was chosen based on ease of
collection and low heavy metal contamination (Allen, 2023). Sediment
was defaunated with a 200
μ
m sieve. The silt/clay-sized fraction of the
sediment was then separated with a 63
μ
m sieve and retained, as par-
ticles larger than this were known to clog lter-feeding structures in the
mussel spat based on previous research (Gui et al., 2016a). Sediment was
thoroughly rinsed with freshwater to remove any soluble fraction that
could interfere with results. To retain the natural characteristics of the
sediment, it was not further sterilised and was refrigerated until use.
To determine reliable concentrations of suspended sediment in
seawater, the relationships among the mass of dried sediment in sus-
pension, the mass of wet sediment in suspension, and turbidity mea-
surements were established for nine suspended sediment concentrations
between 0 and 700 mg L
−1
. The nine concentrations were created by
adding different masses of wet sediment (ThermoFisher Scientic
Sartorius LE2445 analytic scale) from the stock sediment source to 500
mL of FSW in lidded 1 L Schott bottles. Sediments were maintained in
suspension by vigorous shaking, and the bottles were then placed on a
stir plate with a magnetic stir bar inside the bottle for 10 min. The
suspended sediment turbidity was determined by three replicate read-
ings with a YSI ProDSS Swap turbidity meter (Xylem, Yellow Springs,
Ohio). To determine the dry sediment mass of each solution, the bottles
were re-inverted, and three replicate 25 mL samples were ltered with a
Terumo 50 mL syringe through a dried Whatman binder-free glass
microbre lter (25 mm diameter, 0.7
μ
m particle retention). The sy-
ringes were rinsed of remaining particles and salt with 10 mL of Milli-Q
water. Filters were oven-dried at 60 ◦C for 48 h and re-weighed. The
line-of-best-t equation was determined by calibration curves for the
relationships among the corresponding sediment dry weight of the so-
lutions with sediment wet weight and turbidity measurements (Sup-
plementary Fig. S1).
B.S. Biggar et al. Journal of Experimental Marine Biology and Ecology 582 (2025) 152074
2
2.4. Acute SS exposure experiment
Acute effects of suspended sediment on mussel spat were determined
in a 5 day experiment conducted at a range of SS exposure levels. A
series of ten exponentially increasing SS concentrations (0, 5, 10, 25, 50,
100, 250, 500, 1000, 1250 mg L
−1
dry mass sediment) was used for a
dose–response experiment with the <63
μ
m wet-sieved sediment in 1.5
L conical tanks (Supplementary Table S1). To maintain dissolved oxygen
and keep sediment homogeneously suspended, an airstone was sealed
into the base of the conical tank and connected to an aquarium air pump.
A 10 ×15 cm plastic mesh strip (1 mm mesh size) was suspended in each
tank by nylon thread as an attachment substrate for the mussels (Sup-
plementary Fig. S2).
For each tank, 50 green-lipped mussel spat (0.39–4.37 mm SL; mean
1.91 mm) were haphazardly selected from the stock supply, photo-
graphed to determine SL at the outset (using image analyses), and added
freely to the water column of a randomly selected 1.5 L tank.
The wet sediment mass required for each SS treatment was calcu-
lated based on the calibration curve (Supplementary Fig. S1). Wet
sediment was weighed and mixed thoroughly into 100 mL FSW before
being added to the conical tanks holding 900 mL of FSW. After the SS
was added, turbidity was measured with a YSI ProDSS Swap meter
(Xylem Inc., Yellow Springs, Ohio) and again 1 h later to ensure sedi-
ments remained in suspension. Temperature and salinity were measured
twice daily in each tank with a Professional Plus YSI Model 30 (Xylem
Inc., Yellow Springs, Ohio), as was oxygen with a Hach HQ40d Lumi-
nescent Dissolved Oxygen Probe Model LDO101.
The FSW was changed daily by gently pouring tank contents over a
200
μ
m sieve to retain unattached spat. The tanks and mussels were
gently rinsed with freshwater to reduce surface bacteria without dis-
lodging attached mussels. Then, loose mussels were returned to the tank,
followed by fresh sediment solution (as described above) and the
cultured microalgae diet. The mussels were fed a rotation of four
cultured microalgae species, as described above.
At the end of the 5 days, the mussel attachment location in each tank
was noted (i.e., either on the mesh or the tank bottom) and recorded.
Spat were then carefully removed from the tanks, and every mussel was
determined to be living or dead, counted under a dissection microscope,
and photographed to obtain experiment end SL measurements using
ImageJ Software (Rasband, 2011). Spat were considered dead if the
shells were visibly empty or there was no sign of foot, gill, or valve
(opening) movement for 30 min. Some mussels were accidentally lost
during water changes, so the total nal number of mussels per tank was
lower than the initial number. Percent mortality was calculated for each
tank as the number of mussels that were veried dead divided by the
total nal remaining mussels (dead +alive) ×100 %.
2.5. Chronic SS exposure experiment
To determine the chronic effects of SS on mussel spat, a 30 day
experiment was conducted at ve exposure levels of dry sediment mass
(0, 10, 100, 1000, 1250 mg L
−1
), each with ve replicates and 1 g (~
1300 individuals) of green-lipped mussel spat per tank. Based on the
results of the previous experiment, eight treatments within 0–1000 mg
L
−1
were deemed unnecessary, so fewer concentrations were utilised
while maintaining the same overall range (0–1250 mg L
−1
). The tanks,
suspended sediment procedure, daily maintenance, feeding, and phys-
ical measurements were the same as described above for the acute SS
exposure experiment. However, turbidity was not measured 1 h after
adding SS, as it was deemed unnecessary based on the previous exper-
iment. Additionally, to analyse sediment characteristics at the end of the
experiment, the water was collected from daily water changes, retained,
and decanted to recover the sediment (“recovered sediment”from all
treatments pooled).
The number of mussels in 1.0 g (wet weight) was determined before
the experiment by counting the number of spat in ten 0.1 g samples and
then calculating the mean number per gram (i.e., 1329 ±104 (SE) in-
dividuals g
−1
). The average number of individuals per tank was only
calculated as a reference; it was not used in any statistical analyses.
The green-lipped mussel spat (0.31–2.71 mm SL; mean 1.35 mm)
were haphazardly selected from the holding tank and weighed to 1.0 g.
Three randomly selected sub-samples of 50 mussels per tank were
photographed to determine the mean SL at outset. Then, the entire 1.0 g
of mussels was added freely to the water column of one randomly
selected tank. This process was repeated for each tank.
At the end of 30 days, the location of nal spat attachment was
recorded and photographed, and then they were carefully removed from
the tanks. Every mussel was sorted into living and dead (as described
above) under a dissection microscope, counted to determine mortality,
and photographed to obtain experiment end SL measurements. After-
ward, spat were rinsed in freshwater and frozen until further processing.
The experiment end SL of all mussels (i.e., both alive and dead) were
measured from photographs using ImageJ Software (Rasband, 2011).
Spat dry weight was determined by lyophilizing (Christ Alpha 2–4
LSC, Buch &Holm A/S, Herlev, Denmark) all frozen spat for 24 h. The
ash-free dry weight (AFDW) of spat was determined by ashing three
~0.1 g samples per tank of lyophilized mussels (n=75 samples: 3
replicates ×5 tanks per treatment level ×5 treatment levels) for 5 h at
450 ◦C (Nabertherm LT15/11/B410 mufe furnace, Germany). The
total organic matter (TOM, %) of mussel samples was calculated as
(AFDW/total dry weight of sample) ×100 %.
The caloric content per gram of dry mussel was determined using
three ~0.2 g samples per tank of lyophilized spat (n =75 samples: 3
replicates ×5 tanks per treatment level ×5 treatment levels) (Parr 6725
semimicro calorimeter, Parr Instrument Company, USA). This was
converted to caloric content per gram of organic matter using the
percent TOM calculated above for each SS treatment (Supono et al.,
2020). Caloric content, AFDW, and TOM were determined using a
mixture of mussels from different settlement locations within each tank.
The minimum soft tissue requirements to measure caloric content were
too high to allow the separate analysis of mussels per settlement
location.
2.6. Sediment analyses
The AFDW of stock and recovered sediments from the chronic SS
experiment was determined by lyophilizing nine replicate 0.1 g samples
of each for 24 h, weighing, and then ashing in a mufe furnace at 450 ◦C
for 5 h. The total organic matter (TOM, %) of the sediment samples was
then calculated as (AFDW/total dry weight of sample) ×100 %. The
grain size distribution of stock and recovered sediments was determined
by adding 40 mL hydrogen peroxide to 15 mL of sediment for 7 days to
dissolve the organic material. Then, samples were centrifuged and
rinsed with deionised water three times, and 10 mL Calgon was added to
each sample to break up particle aggregates before processing with a
Malvern Mastersizer 3000 (ATA Scientic).
2.7. Statistical analyses
All data analyses and presentations were conducted with RStudio
v4.3.0 (Posit Team, 2023). In all cases, p-values <0.05 were considered
statistically signicant. The two experiments (short and long term) were
analysed separately using linear mixed-effects models (lme4::lmer; Bates
et al., 2015) and generalised linear mixed-effects models (lme4::glmer).
Where models were signicant, contrasts were computed with the
estimated marginal means package (emmeans; Searle et al., 1980) for
factor-level comparisons.
Mortality analyses used a generalised model to specify the family as
binomial (0 =dead, 1 =alive), and tank was included as a random
intercept effect to account for the non-independence of mussels within a
tank. Settlement location was analysed similarly, with a binomial glmer
(0 =bottom of tank, 1 =mesh), and tank was included as a random
B.S. Biggar et al. Journal of Experimental Marine Biology and Ecology 582 (2025) 152074
3
effect.
Mussel growth analyses used an lmer model. All of the measured
mussel lengths were included: for the short term experiment, n=946
(~50 mussels ×10 tanks ×2 time points); for the long-term experiment,
n=29,268 (~250 mussels ×25 tanks at outset and ~ 1000 mussels ×25
tanks at end). To account for the non-independence of mussels within a
tank, a random intercept effect was included. Time (experiment outset
vs. experiment end) was included as an explanatory xed effect to
explore the difference in mussel SL through time. Growth was also tested
between mussel settlement locations (long-term experiment only), using
the same model structure with settlement location included as a xed
effect. Only the SL data from the experiment end were used for this
analysis, so time was not included. As above, tank was included as a
random effect.
The lmer model was also used to test for differences in TOM and
caloric content. Time was not included, as the TOM/caloric content
data from experiment outset were not sufcient for comparison. The
models for the above analyses, as coded in r, are shown below.
glmer(Mortality [or Settlement] ~ SS_Treatment +(1 | Tank), fam-
ily =Binomial, data =df)
lmer(Length ~ SS_Treatment*Time +(1 | Tank), data =df)
lmer(Length ~ SS_Treatment*Settlement_Location +(1| Tank), data
=df)
lmer(TOM [or Caloric_Content] ~ SS_Treatment +(1 | Tank), data
=df)
Differences in physical parameters (temperature, oxygen, salinity)
among tanks were tested separately with two-way repeated-measures
ANOVA to determine whether any differences may have impacted the
results. None of the physical parameters had signicant differences
among tanks for the short-term experiment, so they were not expected to
impact the results and were not included in the nal explanatory models
(Supplementary Fig. S3). Temperature and oxygen were not signi-
cantly different among tanks in the long-term experiment and did not
impact the results of models, and therefore, were not included in the
nal explanatory models for the long-term experiment (Supplementary
Figs. S4, S5). Salinity was signicantly different among tanks, so it was
initially included in the model, however, it had no impact on the model
result and was subsequently removed (Supplementary Fig. S6).
Statistical analyses were not possible for the sediment analyses due
to the way the recovered sediments were collected (all SS treatment
levels pooled, no controls).
3. Results
3.1. Acute SS exposure experiment
Mussel spat mortality was relatively low for all treatments (range =
4–13 %; mean =8 %) and was not affected by SS treatment (p=0.668;
Fig. 1A, Table S2).
Mean mussel spat shell lengths (SLs) were 0.39–4.37 mm (mean =
1.91 mm) at the outset and 0.96–4.56 mm (mean =2.06 mm) at the end
of the acute exposure experiment. In the linear model, SL was related to
time (outset versus end; p<0.001) and the interaction between time and
SS treatment (p=0.03; Table 1). Treatment level comparisons showed
that mussels in 0, 25, and 1250 mg L
−1
had signicantly greater mean
SLs at the experiment end (Fig. 2A, Table S3). Mussels grew over time,
and the greatest growth was in the highest, lowest, and control
treatments.
Mussel attachment location at experiment end, expressed as the
percent of mussels attached to mesh versus the tank bottom, ranged
from 29 % in the 10 mg L
−1
treatment to 87 % in 500 mg L
−1
(mean =
63 %). Mussel attachment to suspended mesh was signicantly related
to SS concentration (p<0.001; Fig. 3A, Table S4), with the lower
concentrations (10, 25, 50, and 100 mg L
−1
) negatively related to mesh
settlement (Table S5). Mussels preferentially settled on the bottom of the
tank in low concentration SS treatments.
3.2. Chronic SS exposure experiment
Mean mussel mortality was relatively low in all treatments (range =
1–30 %; mean =12 %) and not affected by SS treatment (p=0.42;
Fig. 1B, Table S6).
Fig. 1. Percent of juvenile mussel spat (Perna canaliculus) that died following exposure to different levels of suspended sediment concentrations over (A) 5 day short
term and (B) 30 day long term experiments. Grey dots in (B) are the mortality in each tank, and black dots are the mean (±SE) mortality per treatment level. There
was no signicant difference among treatments for either experiment (A or B).
Table 1
Type III ANOVA table summarising the results of linear mixed model to test the
effect of 10 suspended sediment concentrations (SS Treatment) and time
(experiment outset vs. end) on mussel shell length in short term experiment (5
days). * refers to model interaction.
Model Parameter SumSq MeanSq Df F-Value p-Value
SS Treatment 0.81 0.09 9, 2 0.30 0.92
Time 5.27 5.27 1, 924 17.34 <0.001
SS Treatment*Time 5.61 0.62 9, 924 2.054 0.03
B.S. Biggar et al. Journal of Experimental Marine Biology and Ecology 582 (2025) 152074
4
Mussel SLs at the experiment outset ranged from 0.31 to 2.71 mm
(mean =1.35 mm) and 0.61 to 6.55 mm (mean =2.26 mm) at the end of
the chronic exposure experiment. Mean mussel SL was related to both
time (outset vs. end; p<0.001) and the interaction between time and SS
treatment (p<0.001, Table 2). Post hoc contrasts showed that mean
mussel SLs were signicantly greater at experiment end than outset at all
treatment levels (p<0.001; Table S7) and that no treatment levels
differed from each other at experiment outset. The two highest SS
treatments (1000, 1250 mg L
−1
) had signicantly greater SLs at
experiment end than the other treatments (Fig. 2B, Table S8). The
highest SS treatments caused the greatest mussel growth.
Mussel nal attachment location, expressed as the percent attached
to the mesh versus the tank bottom, ranged from 56 % in the 100 mg L
−1
treatment to 83 % at 0 mg L
−1
. Mussel attachment to suspended mesh
was signicantly related to SS concentration (p=0.001; Fig. 3B,
Table S9), with the lower concentrations (10, 100 mg L
−1
) negatively
related to settlement on the mesh (Table S10). Settlement was signi-
cantly higher on the mesh than on the bottom at every treatment con-
centration except for 10 mg L
−1
(p<0.001; Table S11). Settlement
location and its interaction with SS treatment increased mussel growth
(p<0.001, Table 3). Growth was higher on the mesh than the bottom
and in the higher SS concentrations (relative to lower concentrations;
Table S12).
The percent TOM of mussels was 17.8–37.6 % (mean =26.0 %) at
the end of the experiment (Fig. 4A). The caloric content of mussels at
the end of the experiment ranged from 74.3 to 1770.0 cal g
−1
of AFDW
(Fig. 4B). There was no difference in mussel percent TOM (p=0.74,
Table S13) or mussel caloric content among SS treatments (p=0.27,
Fig. 2. Change in mean juvenile mussel (Perna canaliculus) shell length over time exposed to a range of suspended sediment treatments in (A) 5 day short term
experiment and (B) 30 day long term experiment. * represents a signicant change in length over experiment duration (p<0.001 in both experiments, A and B).
Different letters in (B) represent signicant differences among treatments.
Fig. 3. Percent of juvenile mussels (Perna canaliculus) attached to mesh relative to tank bottom following (A) 5 day short term experiment and (B) 30 day long
term experiment.
Table 2
Type III ANOVA table summarising the results of linear mixed model testing
effect of ve suspended sediment concentrations (SS Treatment) and time
(experiment outset vs. end) on mussel shell lengths in long term experiment (30
days). * refers to interaction term.
Model Parameter SumSq MeanSq Df F-Value p-Value
SS Treatment 0.30 0.08 4, 20 0.196 0.94
Time 3052.14 3052.14 1, 29,248 7885.386 <0.001
SS Treatment*Time 13.46 3.37 4, 29,247 8.695 <0.001
B.S. Biggar et al. Journal of Experimental Marine Biology and Ecology 582 (2025) 152074
5
Table S14).
3.3. Sediment analyses
The percent TOM of the sediment was 8.0 ±0.3 % before the
experiment and 4.0 ±2.0 % after. The grain size of the sediment pri-
marily comprised medium and coarse silt (i.e., 15.6–31, 31–63
μ
m)
grain sizes, with 22.8 % of the sample volume in each category. In the
recovered sediment, the grain size distribution was centred on the coarse
silt (31–63
μ
m) size with 51.7 % (Fig. 5).
Table 3
Type III ANOVA table summarising the results of linear mixed model on the
effect of ve different suspended sediment concentrations (SS Treatment) and
mussel settlement location (Location) on mussel growth in long term experiment
(30 days). * refers to interaction.
Model Parameter SumSq MeanSq Df F-Value p-Value
SS Treatment 0.43 0.11 4, 20 0.26 0.90
Location 112.64 112.64 1, 21,022 278.69 <0.001
SS Treatment*Location 45.09 11.27 4, 21,022 27.89 <0.001
Fig. 4. (A) Mean percent total organic matter (TOM) and (B) caloric content of juvenile mussels (Perna canaliculus) exposed to suspended sediment (SS) over 30 day
long term experiment. Grey dots show replicate measurements from experimental tanks, and black dots show mean (±SE) per SS treatment level. There was no
signicant difference among treatments for either measure. AFDW, ash-free dry weight.
Fig. 5. Characteristics of the sediment used in experiments. (A) Mean percent total organic matter (TOM) of sediment (±SE) used in the suspended sediment
treatments in the 30 day long term exposure experiment at the outset of the experiment and as recovered at the end (pooled treatments). Grey dots show replicate
percent TOM samples and black dots are the mean per time period. (B) Grain size analysis of sediment.
B.S. Biggar et al. Journal of Experimental Marine Biology and Ecology 582 (2025) 152074
6
4. Discussion
4.1. Mussel mortality
Small P. canaliculus (<6 mm shell length (SL)) are prone to gill
lament blockage and damage when exposed to suspended food parti-
cles >15
μ
m (Gui et al., 2016b). Hence, exposure to high concentrations
of <63
μ
m suspended sediment could be expected to decrease survival
due to gill damage and blockage. However, the overall mortality of
mussel spat exposed to SS, even at extremely high SS (i.e., 1000), was
relatively low and not different from spat in the treatment without
sediment following the acute (5 day) experiment. This result could have
been due to the short experimental duration. However, at the end of the
chronic exposure experiment (30 day), the mean mortality of mussel
spat was only slightly higher than after the acute exposure experiment,
again with no difference detected among treatments. These results
suggest that green-lipped mussel spat can survive at least 30 days in the
high SS conditions tested in these experiments.
The highest SS concentration tested in the chronic exposure experi-
ment (1250 mg L
−1
) could be expected in highly human-disturbed wa-
tersheds (e.g., agriculture, construction) and/or following an extreme
weather event (e.g., cyclone, ood, landslide) (Ellis et al., 2002;Yanai
et al., 2006). However, these elevated SS levels do not typically last for
≥30 days in coastal oceans and estuaries. Particle settling times in the
ocean are complex and challenging to predict, being inuenced by many
factors, including wind, waves, tides, and particle shape and size (Nasiha
and Shanmugam, 2018;Osborne and Greenwood, 1993;Sutherland
et al., 2015). However, particle settling is substantially increased in
seawater vs. freshwater and high particle concentrations due to aggre-
gates forming with increased density (Kranck, 1980;Lick et al., 1993;
Sutherland et al., 2015). Hence, sediment particles typically settle
through the mixed layer within days (Lande and Wood, 1987), and
prolonged exposure to suspended sediment concentrations of this
magnitude would likely require repeated mass sediment inputs, most
likely the result of cumulative events. For example, in Fiji, following
mass forest burning, a major rainstorm event triggered 620 landslides in
the following weeks (Nunn, 1990). In New Zealand, Cyclone Bola (1988)
triggered 184 landslides per km
2
, and an extreme rainfall event in 2004
triggered thousands of landslides over 16,000 km
2
(Crozier, 2005).
While the results of the current study may not extend to the extreme
events listed here, they do suggest that green-lipped mussel spat survival
is reasonably robust to suspended sediment.
4.2. Mussel shell growth
Mussel shell growth was positively correlated with the interaction
between time and suspended sediment (SS) concentration at both time
scales (5 and 30 days). This result is consistent with previous studies that
have found increased bivalve growth with high levels of SS (Colden and
Lipcius, 2015;Dekshenieks et al., 1993;Emerson, 1990) and, in
particular, clay addition (Aucoin and Himmelman, 2011;Bricelj and
Malouf, 1984;Gatenby et al., 1996;Soniat et al., 1984;Sornin et al.,
1988). Juvenile Elliptio complanata (unionid mussel) and Mercenaria
mercenaria (clam/quahog) both had increased growth in suspended
sediments (Cyr, 2020;Davis, 1960;Dekshenieks et al., 1993), while
Limnoperna fortunei (freshwater mussel) “D”larval stage size was posi-
tively correlated with the inorganic:organic matter ratio (Eilers et al.,
2011). Additionally, Mytilus edulis fed pure algae alone were unable to
reach the same growth rates achieved with silt-addition diets (Soniat
et al., 1984).
Although sediment-mediated growth enhancement is well supported
in the literature, there is little agreement over the mechanism that
supports this growth. Theories include that the sediment provides a
physical substratum for aiding with mechanical digestion, an enzymatic
digestion enhancer, or a source of micronutrients (Gatenby et al., 1996;
Urban and Langdon, 1984). For example, in diet experiments,
Placopecten magellanicus (Atlantic sea scallop) had very low absorption
of organic matter from sediment but very high nitrogen absorption
(Cranford and Grant, 1990). Additionally, sediment may aid in the
retention of small food particles (Deslous-Paoli, 1985;Sornin et al.,
1988) and play a role in pedal feeding in juvenile bivalves with under-
developed feeding structures (Gatenby et al., 1996;Hua et al., 2013;
Reid et al., 1992). While the mechanism behind this pattern of SS-
enhanced growth may be specic to the species and their environ-
ment, its determination could aid in bivalve restoration and aquaculture
work. For example, increased shell growth due to specic nutrients in
the sediment could lead to the formulation of growth-enhancing dietary
supplements.
4.3. Mussel nutritional condition
While sediment-aided growth is primarily regarded as being due to
dietary supplementation through whatever mechanism, the increased
growth could also be viewed as a strategy to limit vulnerability to
environmental perturbation (Aucoin and Himmelman, 2011). The latter
theory spurred the testing of mussel nutritional condition to determine
whether the increased growth had measurable physiological costs.
Prolonged exposure (30 days) to high SS concentrations (up to 1250 mg
L
−1
) was expected to negatively impact mussel condition due to the
following: 1) higher energy expenditures and reduced feeding capacity
from the physical demand and time spent ltering and sorting, and 2)
loss of energy from mucous and psuedofaeces production. However, the
percent TOM and caloric content of mussels showed no differences
among treatments. Although not signicant, there was a trend of low-
ered mean caloric content in the two highest SS treatments, suggesting
that the food intake or energy expenditure of these mussels may have
been marginally altered (Table S15). However, these results show that,
under the conditions tested here, mussel spat can endure high SS (up to
1250 mg L
−1
for 30 days) with no measurable cost to nutritional
condition.
4.4. Mussel nal attachment location
The attachment location of mussel spat at experiment end was
signicantly related to SS treatment in both experiments (5, 30 days),
with more mussels settled on the tank bottom at low SS concentrations.
Mussels have complex settlement behaviour, which may contain mul-
tiple settlement and drift stages (Bayne, 1964). This may explain the
results observed here. If the conditions on the tank became unsuitable at
higher concentrations (more turbid, burial risk), the mussels might have
re-joined the water column to nd a more favourable habitat. These
results suggest that high SS triggers detachment and migration of mussel
spat, which may be an important factor limiting recruitment in wild
mussel populations exposed to episodic high SS events.
Wild and restored mussel beds in New Zealand have been limited by
recruitment, with high sediment loads implicated as the cause. In this
study, suspended sediment had no impact on mortality or nutritional
condition and a positive effect on growth. However, despite the lack of
evidence for sediment negatively impacting mussels, it still triggered
them to detach and depart the tank bottom in high SS concentrations.
Larval settlement is highly dependent on settlement cues and attach-
ment substrates (Alfaro et al., 2006;Wilcox et al., 2020). Suspended
sediment may, thus, disrupt settlement cues and cover settlement sub-
strates, making them unfavourable for mussel settlement. This study
provides strong support for the theory that suspended sediment in-
terrupts mussel settlement and should be explored further in future
research.
4.5. Sediment analyses
The percentage TOM of the stock sediment (8.0 ±0.3 %) agrees with
previous studies on mudat sediments (Cai et al., 2013;Carneiro et al.,
B.S. Biggar et al. Journal of Experimental Marine Biology and Ecology 582 (2025) 152074
7
2021;Cheng and Chang, 1999;Christie et al., 2000), showing that it was
representative of what would likely be experienced in the natural
environment. The percent TOM of the recovered sediment was lower
than the start (4.0 ±2.0 %), suggesting that some of the organic material
was ingested and assimilated by the mussels or utilised through bacterial
activity. Additionally, the mean grain size of the recovered sediment
appears to have shifted towards larger grains, suggesting that ne grains
were retained (e.g., clogging the gills) or aggregated by mucous during
pseudofaeces production and were unrecovered during the sediment
recovery process.
5. Conclusions
This study explored the survival, growth, and nutritional condition
of mussel (P. canaliculus) spat under a range of suspended sediment (SS)
concentrations at two time scales (5 and 30 days) to determine whether
SS could be an explanatory factor in declining mussel populations and
struggling restoration efforts. The results here provide strong evidence
that mussel spat are relatively robust to high SS concentrations over time
scales relevant to extreme weather events, with high survival and
increased growth. However, high SS concentrations triggered detach-
ment and departure, with mussels presumably attempting to escape
unfavourable conditions. This result may explain the low recruitment
numbers in mussel restoration and should be further explored in future
research. Additionally, this study shows potentially advantageous ef-
fects of high SS levels on mussel growth, which may have restoration
and aquaculture implications.
Funding
This research was supported by the Marine Farming Association and
the Ministry for Primary Industries through the Sustainable Food and
Fibre Futures fund #SFFF-23044.
CRediT authorship contribution statement
Brandy S. Biggar: Writing –review &editing, Writing –original
draft, Visualization, Methodology, Investigation, Formal analysis, Data
curation, Conceptualization. Andrew Jeffs: Writing –review &editing,
Validation, Methodology, Funding acquisition, Conceptualization.
Jenny R. Hillman: Writing –review &editing, Validation, Methodol-
ogy, Funding acquisition, Conceptualization.
Declaration of competing interest
The authors declare that they have no known competing nancial
interests or personal relationships that could have appeared to inuence
the work reported in this paper.
Data availability
The data and code are available on Mendeley doi:
10.17632/v74xmgcwgg.1, link: data.mendeley.com/datasets/v74xmgc
wgg/1.
Acknowledgements
This work is dedicated in memorium to laboratory technician and
friend Wednesday Davis; thank you for all your support, advice, assis-
tance, and laboratory chats. We are also grateful to the University of
Auckland’s School of Biological Sciences seawater facilities and labo-
ratory technician Esther Stuck for her assistance and support during the
experiments. We are very grateful to SpatNZ for supplying and trans-
porting all mussel spat utilised in this work and to Luke Johnston for his
discussions and advice on mussel spat rearing. Furthermore, we grate-
fully acknowledge the statistical consultation and advice provided by
Jessica McLay, Statistical Consultant at the University of Auckland. We
also thank Leigh Marine Laboratory and laboratory technician Maria
Mugica for assistance with the calorimetry and AFDW analyses, the
School of Environment (ENV) ShaRE laboratory facilities, and lab
technician David Wackrow for assistance with the grain size analyses.
Additionally, we are grateful to the McCrae family, the Marine Farming
Association, and the Ministry for Primary Industries for providing the
nancial support that made this work possible.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jembe.2024.152074.
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