Journal of the Marine Invertebrate recruitment on artificial substrates
Biological Association of the
United Kingdom
cambridge.org/mbi
Original Article
Cite this article: Siddik AA, Al-Sofyani AA,
Ba-Akdah MA, Satheesh S (2018). Invertebrate
recruitment on artificial substrates in the Red
Sea: role of substrate type and orientation.
Journal of the Marine Biological Association of
the United Kingdom 1–10. https://doi.org/
10.1017/S0025315418000887
Received: 1 June 2018
Revised: 17 September 2018
Accepted: 18 September 2018
Key words:
Artificial panels; artificial reefs; benthic
ecology; biofouling; coastal ecosystem
functioning; larval settlement; Red Sea
Author for correspondence:
S. Satheesh, E-mail: ssathianeson@kau.edu.sa,
satheesh_s2005@yahoo.co.in
© Marine Biological Association of the United
Kingdom 2018
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https://doi.org/10.1017/S0025315418000887
in the Red Sea: role of substrate type
and orientation
A.A. Siddik, A.A. Al-Sofyani, M.A. Ba-Akdah and S. Satheesh
Department of Marine Biology, Faculty of Marine Sciences, King Abdulaziz University, Jeddah, Saudi Arabia
Abstract
Surface physical properties, hydrodynamics, biochemical cues, orientation and temporal scales
play an important role in invertebrate larval recruitment on artificial substrates. In the present
study, invertebrate recruitment on four different substrates (acrylic, stainless steel, ceramic and
concrete panels) was investigated in two different orientations (vertical and horizontal) in the
central Red Sea. Results showed significant variations in the abundance of benthic inverte-
brates between the different substrates. While barnacles and bivalves preferred panels placed
in vertical positions, the abundance of bryozoans was high on horizontal panels. Artificial
panel submersion season plays a significant role in the recruitment of benthic invertebrates
on surfaces in the Red Sea. In conclusion, this study supports the overall notion that marine
invertebrate recruitment on hard substrates is regulated by a combination of factors which
include substrate type, orientation and submersion season.
Introduction
Most marine invertebrates have a complex life cycle which includes a planktonic larval stage
and juvenile or benthic adult stages (Degnan & Degnan, 2010; Calado & Leal, 2015).
Generally, in the marine environment, hard substrates host sessile invertebrates such as bar-
nacles, mussels, ascidians, bivalves, bryozoa, porifera, cnidaria and annelids (Connell, 1999;
Holloway & Connell, 2002; Perkol-Finkel & Benayahu, 2006). The competent larvae in the
pelagic stage begin to search for suitable substratum, where settlement and metamorphosis
occurs. The larvae of sessile invertebrates are induced by a variety of physical, chemical or bio-
logical cues to settle (Rodriguez et al., 1993). Settlement is a process which includes the search
of substratum using larvae-specific sensory mechanisms (Matsumura & Qian, 2014) before
finally metamorphosing to a benthic juvenile.
Biotic and abiotic factors strongly influence invertebrate larvae during the settlement pro-
cess (Rodriguez et al., 1993; Vermeij & Sandin, 2008; Penin et al., 2011). Initially, habitats are
colonized by bacteria and diatoms (Zardus et al., 2008; Satheesh & Wesley, 2011), which may
alter the substrate properties (Lozano-Cortés & Zapata, 2014). Extracellular polymeric sub-
stances from the microbial biofilm (Camacho-Chab et al., 2016), settlement cues from
surface-associated diatoms (Patil & Anil, 2005) and conspecific cues (Da-anoy et al., 2017)
are reported to induce larval settlement. In addition to biotic factors, the experimental results
from many studies reveal that abiotic factors like surface topography, luminous intensity, prox-
imity, surface orientation, hydrodynamics, size and substrate colour influence invertebrate
assemblages on substrates (Thorson, 1964; Maldonado & Young, 1996; Thomason et al.,
2002; Satheesh & Wesley, 2010; Muhammad et al., 2014; Spagnolo et al., 2014; Whalan
et al., 2015).
Introduction of artificial materials due to increasing urbanization is part of major habitat
modifications in coastal ecosystems throughout the world (Vaz-Pinto et al., 2014).
Anthropogenic structures introduced in the water column become a suitable habitat for inver-
tebrate colonization (Chase et al., 2016). Materials used in construction offer potential hard
substrates for settlement of fouling organisms (Whomersley & Picken, 2003; Yan et al.,
2006; van der Stap et al., 2016) and non-indigenous organisms (Cariton & Geller, 1993;
Bax et al., 2003; Page et al., 2006; Ruiz et al., 2009). Numerous studies report higher abundance
and species richness of non-indigenous species on artificial substrates rather than natural habi-
tats (Glasby et al., 2007; Airoldi & Bulleri, 2011; Dafforn et al., 2012). The effect of an artificial
substratum on benthic community recruitment has been studied for the assessment of macro-
fouling assemblages (Anderson & Underwood, 1994; Satheesh & Wesley, 2008), non-
indigenous species (Bumbeer & Rocha, 2016; Marraffini et al., 2017) and artificial reef con-
struction for the purpose of recreational diving, fishing and coral reef restoration (Baine,
2001; Dupont, 2008; Oh et al., 2008). Considering the role of artificial materials in benthic
community colonization and introduction of non-indigenous species, it is important to
study the effects of substrates and their orientation on larval recruitment.
Invertebrate larval settlement studies have shown variation in the community structure
between horizontal and vertical surfaces (Connell, 1999; Glasby & Connell, 2001). Studies
determining the effect of orientation revealed that macrofouling organisms (tube worms
 2 A.A. Siddik et al.

and barnacles) are influenced by substrate orientation (Glasby &

Connell, 2001). In our study, four different substrates: acrylic,
concrete blocks, ceramic tiles and stainless steel plates, which
mimic the materials used in marine applications (Glasby &
Connell, 2001; Stachiw, 2004; Dobretsov, 2015), were used to
test the invertebrate settlement pattern. The following questions
were addressed in our study: (i) Does the artificial substrate
have an effect on larval recruitment? (ii) Is there any significant
variation in invertebrate larval recruitment between different
orientations? (iii) Is the variation in invertebrate recruitment in
relation to substrate type and orientation consistent across tem-
poral scales in the Red Sea? Since the central Red Sea is ultra-
oligotrophic in nature (Almahasheer et al., 2016), studying inver-
tebrate recruitment on artificial substrates assumed significance,
particularly due to the unique environmental conditions.

Materials and methods

Study area
This study was conducted in Obhur Creek, central Red Sea, 35 km
north of Jeddah, Saudi Arabia (21°42′ 33.52′′ N 39°5′ 45.71′′ E).
Obhur Creek (Figure 1) is about 10 km long with a maximum
width of 1.2 km and depth of 35 m and is connected directly to
Fig. 1. Map showing the Obhur Creek and the study area (station) in the central Red
the Red Sea (El–Rayis & Eid, 1997). The creek is a popular
Sea coast of Saudi Arabia.
place for recreational activities and marinas. The temperature of
Obhur Creek varies between 27.16 and 33.22°C and the salinity
varies between 39.82 and 40.22 psu during winter and summer organisms (mean ± SE) was described for the panel surface area
respectively (AlSaafani et al., 2017). of 225 cm−2 (one side of the panel).

Experimental design Statistical analysis

Treatment panels were prepared using four different types of sub-
strate material: acrylic (colourless, transparent sheet, ISO 9001
grade), concrete blocks (a material used in pontoons or jetties),
ceramic tiles (white polished surface) and stainless steel plates.
Treatment panels (each panel: 15 × 15 cm) were randomly
attached to a PVC raft using a cable tie. Each PVC raft contained
four panels (one panel from each substrate) and eight rafts were
deployed. Of these, panels on four rafts were placed in a horizon-
tal position and the other four rafts in a vertical position
(Figure 2). In brief, the experimental design consists of replicate
treatments (N = 4 for each substrate and position, i.e. one panel
from each raft) in all the four seasons.
The rafts were submerged at a depth of 2 m at Obhur Creek
near King Abdulaziz University Marine Research Laboratory
(KAU marine station). The PVC rafts holding the treatment
panels were hung (in vertical and horizontal positions) using a
nylon rope from the KAU marine station jetty and positioned
at the bottom using sufficient weight (concrete blocks).
Treatment panels were submerged for 4 weeks during winter
(December 2016), autumn (February 2017), spring (mid-April–
mid-May 2017) and summer (July 2017) seasons. Panels were
retrieved from the seawater after 4 weeks of submersion in each
season. Panels were immediately transferred to the laboratory
and stored (maximum storage duration: 24 h) at less than 4°C
in a refrigerator for further analysis (Glasby & Connell, 2001).
Marine benthic invertebrates recruited on the panels were ana-
lysed under a dissecting microscope (Leica DMI 3000B) and the
number of organisms recruited on each test substrate (only one
side) was counted (manually) and recorded (Dobretsov et al.,
2013). For horizontal orientation panels, the upper side was ana-
lysed. Organisms from the panels were scraped carefully and kept
in 95% ethanol for further identification using a marine inverte-
brate field guide (Hibberd & Moore, 2009). The abundance of
Marine invertebrate settlement on four different substrates was
analysed using nested ANOVA (analysis of variance) with sub-
strates (nested within season and orientation), seasons and orien-
tation as factors. Additionally, a two-way ANOSIM (analysis of
similarities) was carried out using seasons and substrates as fac-
tors. ANOSIM was made based on the Bray–Curtis similarity
index with 9999 permutations. Non-metric multidimensional
scaling (n-MDS) analysis was used to visualize the similarity
(Bray–Curtis similarity) in benthic community recruitment on
treatment panels submerged in different seasons. SIMPER (simi-
larity percentage) analysis was also carried out based on the Bray–
Curtis similarity index to understand the percentage contribution
of each invertebrate group to the dissimilarities between treatment
panels. ANOVA was carried out using STATISTICA (data analysis
software system, version 10). All other analyses were conducted
using PAST (Paleontological Statistics Software Package, version
3.2; Hammer et al., 2001).
Results
The abundance of invertebrates such as bryozoans, barnacles,
tubicolous polychaetes (tube worms), bivalves and gastropods
was studied in detail as these organisms were commonly recruited
on panels. Results showed that the recruitment of marine inverte-
brates was high (though it differed with submersion season) on
stainless steel panels followed by acrylic and concrete panels
(Figure 3). The total number of invertebrates recruited on the sub-
strates showed significant variations between substrates, seasons
and orientation (Table 1). The orientation of the panels showed
significant effects for the recruitment of most of the invertebrate
groups in a nested ANOVA (Table 2). Further, n-MDS analysis
indicated the effects of season and substrate types on invertebrate
recruitment (Figure 3). Two-way ANOSIM also indicated a

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https://doi.org/10.1017/S0025315418000887
 Journal of the Marine Biological Association of the United Kingdom 3

Fig. 2. Test panels with PVC raft submerged in the marine waters in (a) horizontal and (b) vertical positions. Test panels were prepared from four different
substrates: acrylic, stainless steel, ceramic and concrete.

Fig. 3. Recruitment pattern of marine invertebrates

on artificial panels of four different materials in the
central Red Sea. (A). Total number (abundance,
mean ± SE, N = 4) of organisms recruited on artifi-
cial panels. (B) n-MDS plot showing the interaction
of season and materials on invertebrate recruitment.

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https://doi.org/10.1017/S0025315418000887
 4 A.A. Siddik et al.

Table 1. Three-way nested ANOVA (analysis of variance) results for total number of invertebrates recruited on artificial panels

Factor Test Value F Effect – df Error – df P

Season Wilks 0.1642 15.6641 15 254.3729 0.0000

Substrate(Season × Orientation) Wilks 0.0026 7.9485 135 458.7848 0.0000
Orientation Wilks 0.6002 12.2529 5 92.0000 0.0000
Season, substrate (nested within season and orientation) and orientation were used as factors. P < 0.05 = significant.

Table 2. Three-way nested ANOVA (analysis of variance) results for the recruitment of marine invertebrates between four different substrates (acrylic, stainless steel,
ceramic and concrete)

Barnacles Tube worms Bivalves Bryozoans Gastropods

Factor df F P F P F P F P F P

Season. 3 31.2706 0.0000 36.7945 0.0000 6.1067 0.0007 15.3781 0.0000 8.0522 0.0000
Substrate(Season × Orientation) 27 13.2095 0.0000 12.0347 0.0000 3.0295 0.0000 11.2996 0.0000 5.4655 0.0000
Orientation 1 13.9237 0.0003 13.5490 0.0003 4.3016 0.0407 24.6152 0.0000 0.2700 0.6045
Error 96
Total 128
Substrate (nested within season and orientation), season and orientation were used as factors for ANOVA. P < 0.05 = significant.

significant variation in the abundance of invertebrates in relation
to test panel submersion season and substrate types (Table 3).
Among the invertebrates, tubicolous polychaetes were the
dominant recruits on all the substrates during the four seasons,
though significant seasonal variations (Table 2) were observed
(Figure 4). The maximum abundance of tube worms on acrylic,
stainless steel, ceramic and concrete panels was 160.5 ± 42.16
(autumn, vertical position), 123.75 ± 19.42 (winter, horizontal
position), 101.5 ± 22 (autumn, horizontal position) and 49.75 ±
8.03 (autumn, horizontal position) individuals 225 cm−2 respect-
ively (Figure 4). Nested ANOVA indicated significant variations
in the recruitment of tube worms in relation to season, substrate
type and orientation (Table 2). Bryozoans were the other import-
ant group found on the panels with a maximum abundance of
18.5 ± 3.84 (summer, horizontal position), 53.75 ± 10.96 (spring,
horizontal position), 3 ± 0.91 (summer, vertical and horizontal
positions) and 5.75 ± 1.6 organisms 225 cm−2 (winter, vertical
position) respectively for acrylic, stainless steel, ceramic and con-
crete panels (Figure 4). Bryozoan recruitment also revealed signifi-
cant variation in relation to season, substrate type and orientation
(Table 2).
Barnacle recruitment revealed a maximum abundance of 7.25
± 1.25 (autumn, vertical position), 67.25 ± 14.82 (autumn, vertical
position), 12.25 ± 2.65 (autumn, horizontal position) and 11.5 ±
4.48 organisms 225 cm−2 (autumn, horizontal position) respect-
ively on acrylic, stainless steel, ceramic and concrete panels
(Figure 5). Nested ANOVA showed significant variations in bar-
nacle recruitment among seasons, substrate types and orientation
(Table 2). Bivalve recruitment was also high on stainless steel
panels with a maximum of 17.75 ± 4.49 individuals 225 cm−2
(spring, vertical position). The maximum abundance of bivalves
on acrylic, ceramic and concrete panels was 6 ± 1.29 (autumn,
horizontal position), 8 ± 3.3 (summer, vertical position) and
9.75 ± 3.25 individuals 225 cm−2 (spring, vertical position)
respectively (Figure 5). Bivalve recruitment also varied signifi-
cantly between the seasons, substrate types and orientation
(Table 2).
The recruitment of gastropods was high on acrylic panels, par-
ticularly during the autumn season on the panels submerged in
the horizontal position (22.75 ± 7.99 individuals 225 cm−2). On
stainless steel (3.25 ± 1.12 summer, vertical position), ceramic
(2.25 ± 0.6 summer, vertical position) and concrete panels (2.75
± 1.13 autumn, vertical position), the abundance of gastropods
was very low (Figure 6). The abundance of gastropods showed sig-
nificant variations between different seasons and substrates
(Table 2). The orientation of the panels did not show a significant
effect on the recruitment of gastropods (Table 2).
SIMPER analysis indicated an overall average dissimilarity of
68.97% on the abundance of invertebrates between substrates
(Table 4). Tube worms topped the list with an average dissimilar-
ity of 37.89% and their total contribution was 55.09%. Bryozoans
showed an average dissimilarity percentage of 10.77 (contribution
15.66%) and for barnacles, the dissimilarity was 9.28% (contribu-
tion 13.55%). Bivalves and gastropods showed an average dissimi-
larity of 6.72 and 4.07% respectively (Table 4).
Discussion
Artificial panels are frequently used by marine ecologists to
test various ecological hypotheses (Keough, 1984; Anderson &
Underwood, 1994; Connell, 1999; Satheesh & Wesley, 2011) due
to their ease of handling and determining marine organism
behaviour under various environmental conditions (Glasby &
Connell, 2001; Satheesh & Wesley, 2011). In this study, treatment
panels of four different materials (acrylic, stainless steel, ceramic
and concrete) were used to understand the recruitment prefer-
ences of marine invertebrates. The influence of environmental
variables on the settlement and development of benthic commu-
nities has been reported by many previous studies, particularly
from the biofouling ecology point of view (Rajagopal, 1997;
Satheesh & Wesley, 2011; Masi et al., 2015). In addition to envir-
onmental factors, the substratum type and submerged position
(orientation) in marine waters may also influence the settlement,
recruitment and development of benthic organisms (Glasby,
1999a, 1999b; Glasby & Connell, 2001; Azevedo et al., 2006;
Tyrrell & Byers, 2007; Walker et al., 2007; Satheesh & Wesley,
2010; Vaz-Pinto et al., 2014). Results of our study indicated a sig-
nificant difference in the abundance of invertebrates among treat-
ments. Overall, barnacles, bryozoans and bivalves preferred to
settle on stainless steel panels throughout the study period.

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 Journal of the Marine Biological Association of the United Kingdom 5

Table 3. Two-way ANOSIM (analysis of similarities) results for the recruitment of invertebrates in relation to substrate type and season

Factor Total organisms Barnacles Tube worms Bivalves Bryozoans Gastropods

R P R P R P R P R P R P

Season 0.4313 0.0001 0.2393 0.0001 0.2544 0.0001 0.1338 0.0005 0.1122 0.001 0.0558 0.023
Substrate 0.3739 0.0001 0.1572 0.0001 0.2341 0.0001 0.1096 0.0012 0.0844 0.005 0.0455 0.044
ANOSIM was made based on Bray–Curtis similarity index with 9999 permutations. P < 0.05 = significant.

Fig. 4. Recruitment (abundance, mean ± SE, N = 4) of tube worms and bryozoans on artificial panels submerged in horizontal and vertical positions. (A) Tube
worms, (B) Bryozoans.

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https://doi.org/10.1017/S0025315418000887
 6 A.A. Siddik et al.

Fig. 5. Recruitment pattern (abundance, mean ± SE, N = 4) of barnacles and bivalves on four different materials submerged in horizontal and vertical positions in
the central Red Sea. (A) Barnacles, (B) Bivalves.

Besides stainless steel, ceramic tiles also showed higher recruit-
ment of marine invertebrates. Further, SIMPER analysis indicated
that tube worms and bryozoans were the major contributors of
dissimilarity between different substrates used in this study.
Results also indicated that the abundance of invertebrates on
panels varied significantly among the seasons. Seasonal variations
in biofouling community development on artificial panels have
been reported from the Red Sea (Salama et al., 2018). Ba-akdah
et al. (2016) also reported significant seasonal variations in
epifaunal assemblages associated with macroalgal beds in the
Saudi Arabian coast of the Red Sea. The influence of season on
recruitment was very much apparent in tube worms with a higher
abundance on acrylic and ceramic panels during autumn and
stainless steel panels in winter. Seasonal variations in benthic
community recruitment may also depend on larval availability,
breeding biology of the invertebrates and water current pattern
of the study area (Young & Gotelli, 1988; Bertness et al., 1992;
Brown & Swearingen, 1998; Satheesh & Wesley, 2011).

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 Journal of the Marine Biological Association of the United Kingdom 7

Fig. 6. Abundance (mean ± SE, N = 4) of gastropods on artificial panels submerged in horizontal and vertical positions.

Table 4. Similarity percentage (SIMPER) analysis on invertebrate recruitment patterns between different substrates

Overall average dissimilarity: 68.79

Taxon Average dissimilarity Contrib. % Cumulative % Mean acrylic Mean stainless Mean ceramic Mean concrete

Tube worms 37.89 55.09 55.09 34.8 46 19.4 32.1

Bryozoans 10.77 15.66 70.74 5.06 11.8 1.91 2.06
Barnacles 9.288 13.5 84.25 0.969 15.3 3.78 2.25
Bivalves 6.762 9.831 94.08 1.78 6.13 2 3.31
Gastropods 4.075 5.924 100 4.5 0.875 0.594 0.781

This study revealed that the settlement behaviour of marine
organisms is specific to particular benthic groups and the obser-
vations obtained for one group cannot be applied to another. For
example, except gastropods, all other invertebrate groups observed
in this study showed a significant difference between horizontal
and vertical orientation panels. Hence, the abundance of inverte-
brate larval recruitment may also depend on orientation. Generally,
more diverse and abundant communities were found on the lower
side of surfaces placed in the horizontal position due to limited
stress from physical and biological factors (Sokolowski et al.,
2017). In this study, lower side surfaces of the panels placed in
the horizontal position were not analysed due to the differences
in surface topography of ceramic tiles (one side polished with the
other side being rough).
Substrate orientation is an important physical factor influen-
cing the settlement of larval forms on surfaces (Glasby, 1999a,
1999b; Walker et al., 2007). The results of our study revealed
that recruitment of benthic invertebrates such as barnacles, poly-
chaetes and bivalves was high on panels placed in a vertical
position. Bryozoan and gastropod recruitment was high on hori-
zontal panels. The observed differences in invertebrate abundance
between vertical and horizontal orientation substrates may be due
to the smothering by sediment on horizontal surfaces (Baynes &
Szmant, 1989; Irving & Connell, 2002). For instance, Walker et al.
(2007) reported a lower abundance of barnacles on horizontal
substrates than vertical substrates due to high sediment cover
on the horizontal surfaces. Azevedo et al. (2006) observed higher
recruitment of barnacles on the upper side of horizontal sur-
faces but reported high mortality due to sedimentation. The
only invertebrate group that showed a significantly higher abun-
dance on the horizontal surface was the bryozoans. Normally,
bryozoans prefer horizontal surfaces, which could increase
their resistance to the stress from environmental factors and pre-
dators or their ability to grow over the sediment layer (Irving &
Connell, 2002; Walker et al., 2007). While an abundance of the
bryozoan Watersipora subtorquata on the shaded side (lower
side) of artificial substrates was reported by Connell (1999), a
higher abundance of bryozoans observed in the present study
on the upper side of the horizontal panels indicates that this
encrusting invertebrate group can also settle on sediment-covered
surfaces.
Generally, colonization of marine invertebrates on hard sub-
strates depends on many features of the substrate such as physical
factors, chemical composition, surface roughness, surface charge
and mechanical properties (Chaudhury et al., 2005; Flemming
et al., 2009; Aldred et al., 2010; Satheesh & Wesley, 2010;
Brzozowska et al., 2017). Surface roughness is cited as one of
the major influencing factors determining the settlement of larvae
on hard substrates (Bers & Wahl, 2004). While many laboratory
studies confirmed that surface topography is important for larval

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 8 A.A. Siddik et al.
settlement (Schumacher et al., 2007), under natural conditions
other factors such as microfouling and environmental factors
overplay the role of surface characters in larval settlement
(Vucko et al., 2013; Brzozowska et al., 2017). The results of this
study indicate that surface roughness may not be an important
factor for larval recruitment of many invertebrate groups. This
observation is based on the concrete panels used in this study.
Concrete is one of the common materials used for the construc-
tion of artificial reefs (Fabi et al., 2011) due to its stability and
suitability to create more complex structures (Spagnolo et al.,
2014). The surface roughness of the concrete panels may also
be high compared with other materials used in this study
(Azevedo et al., 2006). The recruitment of marine invertebrates
on concrete panels was lower than that on stainless steel and
acrylic panels (except tube worms). The possible reason for the
low abundance of invertebrates on concrete panels might be
environmental factors, particularly accumulation of silt in the sur-
face crevices. The accumulation of silt on surfaces may inhibit the
colonization of many benthic organisms (Azevedo et al., 2006).
There are conflicting reports in the literature regarding the
influence of substrate material on benthic community recruit-
ment. While some studies (Connell & Glasby, 1999; Glasby,
1999a, 1999b; Nandakumar et al., 2003; Azevedo et al., 2006)
reported variations in benthic community composition between
different substrates, others (Creed & DePaula, 2007; Burt et al.,
2009; Dobretsov, 2015) observed no difference in benthic com-
munity among the substrates. In this study, results strongly sup-
port the hypothesis that benthic community recruitment varied
between different substrates. It should be noted here that the pre-
sent study focused on the abundance of benthic groups instead of
the diversity of organisms. The abundance of organisms was
indeed higher on stainless steel than on other materials, at least
for barnacles, bryozoans and bivalves. Many of the previous
reports on the difference in benthic communities between sub-
strates suggested that the variations may be due to the orientation
of the substrates, environmental conditions of the material
immersion site and submersion time (Osman, 1977; Qiu et al.,
2003; Field et al., 2007). Although the orientation of the substrates
in this study was found to be significant for most of the inverte-
brate groups, the differences in the abundance among treatments
such as acrylic, stainless steel, ceramic and concrete raise further
questions as to whether the variations are due to substrate prop-
erties or environmental conditions.
Many previous studies (Tamburri et al., 2008; Hadfield, 2011;
Hadfield et al., 2014; Freckelton et al., 2017) suggested that the
microfouling developed on the substrates may induce settlement
of marine invertebrates. Nandakumar et al. (2003) reported con-
siderable variations in microfouling development between three
substrates. Further, microbial community assemblage on artificial
panels has been shown to be influenced by substratum type
(Chung et al., 2010; Witt et al., 2011; Li et al., 2014). Hence, it
is noteworthy to study the site-specific variations, mainly differ-
ences in microfouling development between the materials and
their further role in benthic community settlement.
In conclusion, the results obtained in this study can be sum-
marized as follows: (1) benthic community recruitment in the
central Red Sea showed strong seasonal variations; (2) preference
of a particular substrate and orientation depend on benthic inver-
tebrate taxa, most likely reflecting the settlement behaviour of
marine invertebrates; and (3) significant seasonal variations in
the abundance of invertebrate recruitment underline the role of
site-specific processes along with substrate and orientation.
Above all, this study confirmed that invertebrate colonization
on artificial materials immersed in seawater is controlled by a
combination of factors. Most of the factors may be local environ-
mental conditions and hence further region-specific studies will
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https://doi.org/10.1017/S0025315418000887
improve our understanding of benthic community recruitment
and coastal ecosystem functioning in the Red Sea.
Financial support. This project was funded by the Deanship of Scientific
Research (DSR) at King Abdulaziz University, Jeddah, under grant no.
G-222-150-38. The authors, therefore, acknowledge with thanks DSR for
technical and financial support.
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