Supplemental Information
In this supplement we provide the effect size data used in our analysis as well as our own
assessment of whether particular studies should be classified as “mainly” epifauna or infauna. To
make these assessments, we followed a very similar process to that of Thomsen & Wernberg
(2015). However, in addition to focusing on whether sampling was done under algae or including
the algae, we based our assessments on whether other features of the sampling, the natural history
of species mentioned in the paper, and information from study authors. In some instances, these
differences in approach led to different assessments. Given the difficulties and subjectivity
involved in some if these assessments it is difficult to know which is the ‘correct’ one. However,
the results of both our analysis and that of Thomsen & Wernberg (2015) are broadly similar
where they overlap.
Table S1: Effect sizes (Hedge’s g) and variances used in the analysis of macroalgal blooms and
mats effects on invertebrate abundance. Experimental studies (Expt) are entered in the table in the
same order used by Thomsen & Wernberg (2015), followed by two studies that they excluded,
and then the observational studies (Obs). Notes explain differences between our approach and
that of Thomsen & Wernberg (2015), address related comments in their supplemental material, or
provide other useful information.
Study
Franz & Friedman 2002 Addition
Franz & Friedman 2002 Removal
Gamenick et al. 1996
Lewis et al. 2003
Norkko & Bonsdorff 1996
Österling & Pihl 2001 Exp 1
Österling & Pihl 2001 Exp 2
Rossi 2006 Site 1
Rossi 2006 Site 2
Rossi 2006 Site 3
Sundbäck et al 1996 Exp a
Sundbäck et al 1996 Exp b
Sundbäck et al 1996 Exp c
Sundbäck et al 1996 Exp e
Thomsen & McGlathery 2006
Wetzel et al. 2002
Aarnio & Mattilda 2000 Exp 1
Aarnio & Mattilda 2000 Exp 2
Bolam et al. 2000
Bolam & Fernandez 2002
Deegan et al 2002
Holmquist 1997
Thomsen 2010
Thomsen et al. 2012
Cummins et al. 2004
Lavery et al. 1999 Infauna
Lavery et al. 1999 Epifauna
Bonsdorff 1992
Defew et al. 2002
Jones & Pimm 2006
Kotta & Orav 2001
Lardicci et al. 2001 West Lagoon
Lardicci et al. 2001 East Lagoon
Lardicci et al. 2001 Channel
Migne et al. 2011 Infauna
Norkko & Bonsdorff 1996
Berezina & Golubkov 2008
Berezina et al. 2009 Gulf
Berezina et al. 2009 Estuary
Deegan et al. 2002
Farina et al. 2003
Garcia et al. 2010
Jephson et al. 2008
Kotta et al. 2008
Migne et al. 2011 Epifauna
Norkko et al. 2000
Wennhage & Pihl 2007
Hedge's g
Variance
-1.03
-0.56
-3.69
-1.90
-0.39
-0.74
-2.29
-2.32
0.32
-2.13
-0.78
-0.90
-1.55
-0.40
-0.76
-0.23
7.49
2.09
3.38
4.49
-0.28
10.10
0.72
2.16
-0.99
0.44
-2.28
-1.62
0.77
-0.59
-2.47
-1.98
-9.31
0.46
0.84
-2.46
-2.61
-0.19
-0.34
-1.28
-6.27
-0.12
-0.44
0.51
0.11
1.57
0.30
0.15
0.15
2.39
1.05
0.29
0.41
0.71
0.32
0.19
0.30
0.48
0.49
0.63
0.44
0.04
0.18
4.03
0.43
1.10
1.12
0.20
2.76
0.11
0.15
0.03
0.44
1.58
0.14
0.01
0.07
0.22
0.13
0.13
0.13
0.08
0.63
0.52
0.27
0.29
0.27
3.53
0.23
0.18
0.07
0.07
0.32
0.03
Study
Type
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Expt
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Obs
Infauna/
Epifauna
Infauna
Infauna
Infauna
Epifauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Epifauna
Infauna
Epifauna
Epifauna
Infauna
Infauna
Epifauna
Epifauna
Epifauna
Epifauna
Infauna
Infauna
Epifauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Note
1
1
2
3
4
5
6
7
8
9
10
11
11
12,13
12,14
12,15
12,16
12,17
12,17
12,17
12,18
12,19
12,20
12,21
12,21
12,22
12,23
12,24
12,25
12,26
12,27
12,28
12,29
Table S1 Notes (Supplemental Information from Thomsen and Wernberg 2015 will often provide context):
1. Effects calculated from the mean and SD of the difference between control and ‘treatment’ using temporal
replicates. The high and low effect sizes were then combined via fixed effects meta-analysis to give single
effect.
2. As noted by Thomsen and Wernberg (2015) Fucus is not normally considered a bloom forming species.
Although most of the data we compiled comes from bloom-forming species, our focus was on the effects of
algal blooms and mats, so we also included mats of non-blooming species such as Fucus. Although we
agree that most of the sampled organisms were likely infauna, note that comments about
gastropods/epifauna not being sampled in the Supplemental Information of Thomsen and Wernberg (2015)
are incorrect. At two species (both Hydrobia species) are specifically mentioned and reported on as a major
component of the community. Hydrobia spp. are also one of the major components of the community in
another paper that the authors have classified as studying epifauna. This example and the two below
(comments 3 & 4) highlight the difficulty of classifying studies.
3. Classified by Thomsen and Wernberg (2015) as ‘mainly infauna’ because core was used. However, the core
also sampled the algae and its associated epifauna (personal communication, Lesley Lewis, 2014). Thus,
according to their criteria, this study should be classified as ‘including algae/mostly epifauna’). Also, an
epifaunal species, Hydrobia ulvae, makes up more than 50% of the average abundance in 3 of the 4 values
used to calculate the effect size.
4. Thomsen and Wernberg (2015) classified this study as ‘under algae/infauna’. Although animals on the algae
(epibionts) may not have been sampled, the study appears to focus on effects of blooms on epifauna. It
examines recruitment onto a hard substrate (brick), sediment is considered a ‘stress’, and all of the animals
mentioned in the paper live on the bottom, algae, or hard structures (not in the sediment). Thomsen and
Wernberg (2015) calculated separate effect sizes for replicates located in three different positions relative to
oyster reefs while we calculated a single effect size for all replicates in this experiment. This is consistent
with how we treated other small-scale multifactorial experiments included in this analysis (and our broader
one).
5. Thomsen and Wernberg (2015) classified this study as ‘mainly epifauna’ because of a statement about
epibenthic gammarids increasing but acknowledged that this classification was difficult to make. We choose
to classify it as infauna because the authors refer to their diversity measure as ‘infaunal diversity’ and,
although ‘epibenthic’ gammarids are mentioned, they are clearly a minor component of the community
compared to the infaunal species. Moreover, several measures (including ‘epibenthic gammarids’) are
referred to as quantities ‘under weed mats’, suggesting epifaunal invertebrates in algal mat were not
sampled. Rather, it seems algae may have been removed from cores in the same way as described for the
sediment sample cores, and the gammarids may have been found at the sediment surface under the algae.
6. Thomsen and Wernberg (2015) classified this study as ‘mainly epifauna’ because the algae were sampled in
the cores along with the sediment. We classified this study as “mainly infauna” because it is clear from the
data that the vast majority of the individuals sampled were from infaunal species, and that the increase in
abundance in plots with algae was due to an increase in infauna species (e.g. Pygospio elegans, Capitella
capitata, and Corophium volutator, among others). Moreover, the only epifaunal taxon mentioned in the
manuscript is Doto sp., a nudibranch that occurred only in plots with algae. It was only moderately
abundant.
7. The effect size of Thomsen and Wernberg (2015) was apparently calculated from the macroepifauna data
only. Ours was calculated by combining the effect sizes for macroepifauna and decapods (also epifaunal
invertebrates, but not included in the macroepifauna samples).
8. Thomsen and Wernberg (2015) elected to calculate separate effect sizes for replicates at two different
depths. This is not consistent with how results from this, or other multifactorial experiments, were included
in our analysis. We first calculated an effect size for each level of the secondary (non-bloom) treatment(s)
and then estimated a single effect size for the study using a fixed effects analysis. In addition, they used
only the replicates for the highest algal density, whereas we used data from both algal densities and the
control.
9. As above, Thomsen and Wernberg (2015) elected to calculate separate effect sizes for replicates at three
positions relative to nearby oyster reefs while we calculated a single effect size for all replicates in this
experiment.
10. Thomsen and Wernberg (2015) excluded this study, stating that it reported on results for individual species
(see their supplemental information). This is not the case. It reports on 8 multispecies abundance
measurements and one single species (Potamopyrgus antipodarium) measurement. As explained in our
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
27.
28.
29.
original manuscript, we used any multispecies measurement as a ‘community’ measure. Following our
protocol (Lyons et al. 2012, Lyons et al. 2014), we estimated our single effect size using a fixed effects
analysis of the 8 multispecies measures (i.e. not including the single species measure). Most of the
multispecies groups used in our estimate are composed primarily of infaunal species and, given the
ecosystem where this study occurred, it is likely that the organisms from groups that include epifaunal and
infaunal species (e.g. bivalves) were primarily infauna. In their supplemental materials, Thomsen and
Wernberg (2015) also mention that summing the abundances of the 9 measures suggests that there are more
invertebrates where algae is present, and that this supports the ‘functional matching’ hypothesis. However,
this is potentially misleading for two reasons: first, as the figure caption of Cummins et al. 2004 (Figure 1)
states, the abundance of only selected taxa are presented. So, summing the presented abundances does not
give us a full accounting of invertebrate abundance any more than our decision to leave out the single
species data; second, this apparent pattern is driven almost entirely by Potamopyrgus antipodarium, which
were present in the algae when it was transplanted to create the ‘bloom’ treatment (Cummins et al. 2004).
Although animals can be transported with drifting macroalgal mats, it appears that the increase in this
species in the bloom treatment is largely artifactual.
In our original paper, the effect sizes for epifauna and infauna from Lavery et al. 1999 were combined into a
single estimate using fixed effects meta-analysis. These studies were excluded by Thomsen and Wernberg
(2015) because, by the time of sampling, algal abundance in the ‘bloom’ treatment replicates had declined
to levels similar to (or less than) that found in the ‘control’ treatment replicates. We acknowledge this as a
reasonable decision (algal-dependent epifauna in particular may decline as algal abundance does), but we
chose to include these studies here (and in original analysis) because the treatments were in place for several
months, and blooms’ effects often last even if the bloom has largely rotted away (or been washed away)
because it takes time for animals to recruit or recolonize areas. Exclusion of these studies from our analysis
has little effect on our estimates (Re-estimated Hedge’s g (95% CI); Infauna Observation: -2.07 (-3.15 to 0.99), Infauna Experiment: -0.29 (1.32 to 0.75), Epifauna Observation: -0.43 (-1.31 to 0.46), Epifauna
Experiment: 1.36 (0.19 to 2.5)).
Not included by Thomsen and Wernberg (2015), who examined only experimental studies.
Invertebrates “under algae” sampled by cores.
Tubificoides spp. and other oligochates sampled by cores
“Infaunal abundance” was sampled
Study focused on infauna
Organisms were mainly infauna (personal communication, Claudio Lardicci, 2014)
Meiofauna were sampled from inside the sediment (personal communication, Aline Migne, 2014) and the
community was dominated by an epifaunal gastropod, Hydrobia ulvae.
Animals “under algae” were sampled in cores.
“Benthic animals, from the bottom sediments and from the attached algae, were collected
Sampled by dip nets, with organisms removed from algae, gravel, etc.”
Decapods sampled by throw nets
Everything in the plot counted or assessed for cover
“Both infaunal and epifaunal invertebrates were sampled in each core.”
Zostera cut at base and all animals associated with each Zostera sample counted.
Drift algae and associated animals collected with grabs
Macroafauna both in and under algae sampled (personal communication, Aline Migne, 2014).
Animals collected from the algae.
Drop trap sampling of epifauna. This estimate is for the Mysid community and does not include the data on
fish or other taxa.
Table S2: Effect sizes (Hedge’s g) and variances used in the analysis of macroalgal blooms and
mats effects on invertebrate species richness.
Study
Bolam & Fernandez 2002
Bolam et al. 2000
Cummins et al. 2004
Franz & Friedman 2002 Addition
Franz & Friedman 2002 Removal
Gamenick et al. 2006
Lavery et al. 1999
Norkko & Bonsdorff 1996
Rossi 2006 Site 1
Rossi 2006 Site 2
Rossi 2006 Site 3
Wetzel et al. 2002
Holmquist 1997
Lavery et al. 1999
Lewis et al. 2003
Thomsen & McGlathery 2006
Thomsen et al. 2012
Thomsen 2010
Bonsdorff 1992
Jones & Pinn 2006
Lardicci et al. 2001 West Lagoon
Lardicci et al. 2001 East Lagoon
Lardicci et al. 2001 Channel
Norkko & Bonsdorff 1996
Ouisse et al. 2011
Deegan et al 2002
Hedge's g
Variance
2.65
-0.45
-1.57
-0.30
-0.79
-6.24
0.59
-4.28
-1.13
-1.79
-0.29
-1.07
10.42
-2.07
-1.05
-0.08
0.22
-0.02
-0.80
-1.13
-12.77
-3.23
-2.33
-2.98
1.24
1.10
0.58
0.37
0.23
0.08
0.18
5.80
0.46
1.05
0.22
0.26
0.19
0.20
2.94
1.55
0.72
0.04
0.10
0.10
0.11
0.08
2.39
0.27
0.20
0.77
0.11
0.36
Study
Type
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
experiment
observational
observational
observational
observational
observational
observational
observational
observational
Infauna/
Epifauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Epifauna
Infauna
Infauna
Infauna
Infauna
Infauna
Infauna
Epifauna
Epifauna
Note
1
2
3
4
4
5
6
7
8
8
8
9
10
6
11
12
13
14
15
16
17
17
17
18
19
20
Table S2 Notes:
1. A total of 17 species were observed. In the manuscript, 10 of the species are mentioned and 9 are infaunal
(and it is very likely those lumped together as ‘oligochates’ were as well), indicating that the majority of the
species observed in the study were infaunal.
2. Authors refer to diversity measure as ‘infaunal diversity’ and, although ‘epibenthic’ gammarids are
mentioned, they are clearly a minor component of the community. Moreover, several measures are referred
to as measures made ‘under algal mats’, suggesting epifaunal invertebrates in the algal mat were not
sampled.
3. Although epifaunal amphipods and isopods and one species of epifaunal gastropod are mentioned, the
identity of most species mentioned in the paper, and the taxonomic groups presented in Figure 4, suggest
that the majority of the species found in this study were infaunal.
4. The data we used came from experiments looking at the “infaunal copepod community”.
5. Most species and groups mentioned in the manuscript are infauna.
6. In our original manuscript separate estimates for epifauna and infauna were combined by fixed effects metaanalysis to give a single, overall effect size.
7. Cores taken after mats removed, suggesting that sampled species were mostly infauna. Also, most of the
species or groups mentioned are infauna.
8. “I investigated how small-scale burial of Ulva spp. affected spatial variation of macrofauna in intertidal
sediment.”
9. Focuses on meiobenthic nematodes.
10. Sampled mobile invertebrates associated with algae and seagrass using throw traps.
11. A mix of several epifaunal and several infaunal species were sampled in this study.
12. Thomsen and Wernberg (2015) classified this study as ‘under algae/infauna’. However, it examines
recruitment onto a hard substrate (brick), sediment is considered a ‘stress’, and all of the animals mentioned
13.
14.
15.
16.
17.
18.
19.
20.
in the paper live on the bottom, algae, or hard structures (not in the sediment). Thus we have classified it as
studying epifaunal invertebrates.
Sampling appears to have included animals living on the algae, seagrass, and sediment, as well as infauna
but descriptions suggest that epifaunal inverts were more numerous.
Sampling did not include infauna.
Invertebrates “under algae” sampled by cores.
“Infaunal abundance” was sampled
Organisms were mainly infauna (personal communication, Claudio Lardicci, 2014)
Animals “under algae” were sampled in cores.
Meiofauna were sampled from inside the sediment (personal communication, Aline Migne, 2014) and the
community was dominated by an epifaunal gastropod, Hydrobia ulvae.
Decapods sampled by throw nets
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